LIBRARY OF THE

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AT URBANA-CHAMBMGN

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FI

v.42

cop. 3

NATURAL H1ST0RK

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3

FIELDIANA • ZOOLOGY

Published by
CHICAGO NATURAL HISTORY MUSEUM

Volume 42 October 17, 1958 No. 6

Philippine Zoological Expedition

1946-1947

STONEFLIES FROM THE PHILIPPINES
(PLECOPTERA)

Stanley G. Jewett, Jr.

Among the insects collected by the Philippine Zoological Expe-
dition, 1946-47 (Hoogstraal, 1951), are a number of well-preserved
Plecoptera, nymphs as well as adults. For the most part these were
taken on the island of Mindanao; a few are from Palawan. The
material includes nine species, one of which I have described as new.
I have included descriptions of two nymphs which are of particular
interest because they represent distinct phylogenetic lines apparently
confined to Asia.

A large number of names have been applied to neoperline stone-
flies from southeastern Asia. Many of these may eventually be
synonymized. A detailed study of material from the East Indies
and the mainland of southeastern Asia should be made to determine
the exact number of species; a number of them may occupy a wide
geographic range, including the Philippines. Klapalek (1921), Navas
(1918), and Banks (1913, 1920, 1924, 1937) have described neoper-
lines from the Philippines. While it has not been possible for me to
examine the types of species described from the Philippines I feel
that it is reasonably correct to associate some of Banks' names with
material contained in this collection.

I have followed Kimmins (1950) in regarding Phanoperla Banks
(1938) as a genus distinct from Neoperla, because of the distinctive
venation and male genitalia. I regard Rhopalopsole Klapalek (1912)
as a subgenus of Leudra, since it has venation almost identical with
the subgenus Paraleuctra Hanson (1941), and its prosternal sclerotiza-
tion is like that of the subgenus Zealeuctra Ricker (1952) ; its peculiar

Library of Congress Catalog Card Number: 58-59918 rr T T

No. 857 77 r\ /f "V*

NATURAL it"",TSTv T P-lTOS

HISTORY SURVEY

78 FIELDIANA: ZOOLOGY, VOLUME 42

male genitalia furnish sufficient reason for regarding it as a distinct
subgenus.

The material listed in this paper is in the collection of Chicago
Natural History Museum unless otherwise designated, as follows:
USNM (United States National Museum), MCZ (Museum of Com-
parative Zoology), and SGJ (the writer).

I am indebted to Dr. P. J. Darlington, Jr., Museum of Compara-
tive Zoology, through whose courtesy I was able to examine two
specimens of Leuctra (Rhopalopsole) malayana Banks, including the
male type; to Dr. J. F. Gates Clarke, United States National Mu-
seum, for the loan of the Philippine stonefly material under his care,
most of which was studied by Banks; and to Dr. Clifford C. Gregg,
Director of Chicago Natural History Museum.

Family PELTOPERLIDAE

Peltoperla mindanensis Banks. Figure 17.

Peltoperla mindanensis Banks, 1924, Bull. Mus. Comp. Zool., 65: 426, pi. 2,
fig. 20.

The male genitalia of this species (fig. 17, A) are notable for the
median lobe on the anterior median border of the ninth sternite.
The female subgenital plate (not figured), a large produced lobe ex-
tending over the ninth sternite and evenly rounded, is typical for
the genus. Teneral specimens have remnants of the thoracic gills,
but these are absent in fully hardened adults.

Nymph (nearly mature) : Length of body 10 mm. Body brown, covered with
fine, appressed hairs, somewhat less roach-like in general aspect than Peltoperla
brevis Banks. Single, slender, finger- like external gills present as follows: under
each side of sternal plates of prothorax and mesothorax at base of legs, under each
side of dorsal plates of mesothorax and metathorax, and on each subanal lobe (fig.
17, C). Stout spines or bristles distributed as follows: as a border on margins of
sternal plates, that of anterior margin of prosternal plate longest and most con-
spicuous; as a fringe around hind margin of each abdominal segment and each
cereal segment; on subanal lobes; on outer margin of coxae; scattered on outer
face of femur and as a terminal band; scattered on tibia and as a terminal band.
There is a row of fine, long hairs on outer margins of each femur and tibia, those
on tibia longest. Each femur deeply grooved below to hold tibia when folded;
each femur and tibia compressed vertically to lie close to sides of thorax. Each
sternum of thorax with a plate overlapping backward (fig. 17, B). Cereal seg-
ments normal, well over 25 in number.

Galea reaching lacinia, with a tuft of hairs at the tip (fig. 17, D); lacinia with
two spines at tip and an inner spine; paraglossae exceeding glossae, both with tuft
of hairs at tip; mandibles with several wedge-shaped teeth.

^ (,

JEWETT: STONEFLIES FROM THE PHILIPPINES

79

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.3

Until more is known about the nymphal morphology of Asiatic
peltoperlines, it seems best not to attempt to assign this species to
a subgenus.

Material examined. — Mindanao: Todaya, Mount Apo (east
slope; 2,800 ft. alt.) ; 22 N, 2 <? , 5 9 (10 N, 1 d\ 2 9 , SGJ) ; on rocks
in and along the Sibulan River; Oct. 27-Nov. 20, 1946; D. Heyne-
man and G. Alcasid.

A B D

Fig. 17. Peltoperla mindanensis: A, male genitalia, ventral view; B, nymphal
sternal plates; C, right anal gill of nymph; D, left maxilla of nymph.

Camp Meran, Mount Apo (east slope) ; 1 9 ; "along a swift
stream;" Nov. 6, 1946; H. Hoogstraal.

Mount McKinley (east slope; 3,000-3,300 ft. alt.); 1 N (small),
1 9 ; on rocks in creek and along stream; Aug. 12-29, 1946; H. Hoog-
straal and F. G. Werner.

Bugasan-Parang, Cotabato Province; 1 d" (SGJ); at light; Dec.
12, 1946; F. G. Werner.

Surigao; 1 <? (USNM); C. F. Baker.

Family NEMOURIDAE
Subfamily Leuctrinae
Leuctra (Rhopalopsole) malayana Banks. Figure 18.

Leuctra malayana Banks, 1920, Bull. Mus. Comp. Zool., 64, (3), p. 325, pi. 4,
fig. 45.

80 FIELDIANA: ZOOLOGY, VOLUME 42

The following descriptions are based on material from Mindanao.
The male type of this species and a female bearing an identical local-
ity label, "Imugin, N. Viscaya," were kindly lent to me. These two
pinned specimens were partially relaxed, examined, and found to be
virtually identical with the Mindanao material. The furcate tips of
the lateral prongs of the male genitalia in the type are more promi-
nent than in the male from Mindanao but are similarly shaped; the
female from Imugin has a subgenital plate that appears to be iden-
tical with that of the Mindanao females.

Nymph (mature): Length of body 8.5 mm. Entire body (fig. 18, C) yellow
brown, mature wing pads nearly black. Head rounded; antennae slender, straight,
without hairs. Ocelli translucent, almost invisible. Dark marks present in front
of anterior ocellus (absent in immature nymphs).

Presternum and basisternum separated by a groove. Entire body without
hairs. Each cereal segment with a distal whorl of long bristles.

Galea exceeding lacinia (fig. 18, D), without hairs; lacinia with fringes of hairs
on both upper and lower margins; labium typical; mandibles with several heavy
teeth and basal to these a blade whose outer half is margined by a row of stiff
setae (fig. 18, E).

Adult: Length to wing tips: male 10 mm., female 8.5-12 mm. Length of body:
male 8 mm., female 7-10 mm. Body brown, wings slightly fumose. Venation
typical for subgenus, with no fork in Cu! of hind wing beyond vein cu-m. Pre-
sternum completely separated from basisternum.

Male: Dorsal processes absent from abdominal segments. Ninth and tenth
segments and their appendages heavily sclerotized; a median area on ninth tergite
somewhat membranous. Other abdominal segments largely membranous, with
sclerotized areas on anterior borders of the tergites except in the wholly mem-
branous median quarter of each. Anterior border of ninth sternite bears a hairy
median lobe, beneath which is a clear area (fig. 18, B); the lobe protrudes out from
the sternite at an angle of about 45 degrees. The ventral distal sclerite on the ninth
sternite ends in a rounded, median lobe which is membranous at its tip. The pos-
terior median portion of the tenth tergite is formed into a small, recurved, sclero-
tized, dorsal hook (fig. 18, A). Lateral portions of the tenth tergite are greatly
enlarged and drawn out distally to form furcate prongs. Cerci beset with hairs;
cerci arise from lateral base of tenth tergite. Hairs on eighth sternite longer and
more dense than those on seventh and ninth sternites.

Female: Subgenital plate on seventh sternite slightly produced, rounded, trans-
lucent medially at its tip (fig. 18, F); what appears to be a rounded notch at its
tip is the sclerotized genital opening. Eighth sternite unmodified.

Material examined. — Mindanao : Lake Linau, Mount Apo (north
slope; 7,900 ft. alt.) ; 16 N, 1 tf1 , 3 9 (6 N, 1 cf, 1 9 , SGJ) ; in mossy
forest, and in flight over stream through tall grass; Oct. 25-Nov. 2,
1946; H. Hoogstraal and F. G. Werner.

Burungkot, Upi, Cotabato Province (1,500 ft. alt.) ; 1 9 (SGJ) ;
at light; Jan. 1, 1947; F. G. Werner.

JEWETT: STONEFLIES FROM THE PHILIPPINES 81

B

:\J-

.\j.

Fig. 18. Leuctra (Rhopalopsole) malayana: A, male genitalia, lateral view;
B, male genitalia, ventral view; C, outline of nymph; D, left maxilla of nymph;
E, left mandihle of nymph; F, female genitalia, ventral view.

Crater Lake, Mount Apo (9,000 ft. alt.); 1 9 ; Nov. 15, 1946;
H. Hoogstraal.

Luzon: Imugin, northern Viscaya; 1 d" (type, MCZ), 1 9
(MCZ);C. F. Baker.

Leuctra (Rhopalopsole) palawana, new species. Figure 19.

Holotype. — A female, south slope of Mount Balabag, Mantalin-
gajan Range, southern Palawan Island, Philippine Islands, May 4,
1947; F. G. Werner. In the collection of Chicago Natural History
Museum.

The shape of the subgenital plate and the sclerotized areas on
the eighth sternite serve to distinguish this species from L. mala-

82 FIELDIANA: ZOOLOGY, VOLUME 42

yana Banks, the only other described species of Leuctra from the
Philippine Islands.

Female: Length to tip of wings 8.5 mm., to end of body 7 mm. In general
appearance very similar to L. malayana, described above. The distal portion of
the subgenital plate is produced and slightly rounded; eighth sternite sclerotized
as in figure 19.

Family PERLIDAE
Subfamily Perlinae
Genus Neoperla Needham

Pseudoperla Banks, 1892, Trans. Am. Ent. Soc, 19: 322 (not Pictet, 1854,

Traite Pal., 2, ed. 2, p. 364).
Neoperla Needham, 1905, Proc. Biol. Soc. Washington, 18: 108.

The separation of Asiatic species in this genus is very difficult
because of wide variation in morphological characters in many of
the species and because most of the present descriptions do not in-
clude sufficient details, particularly of male genitalia, to separate
them. While the size and location of the eyes and ocelli are quite
variable, these structures aid in separating the species when corre-
lated with general head shape and the morphology of the rear median
area of the head. It is particularly desirable to preserve fresh mate-
rial of this genus in alcohol to preserve color patterns and to avoid
the distortion which occurs when specimens are dried.

Four distinct species are present in the material taken by the
expedition. Additional material from Mindanao and Luzon (bor-
rowed from the United States National Museum) includes the same
four species. It is of interest to note that two of these species occur
on both islands. It is quite possible that some or all of the species
occur in parts of southeastern Asia. The male genitalia of the
four species closely resemble those of Neoperla clymene (Newman),
the type species of the genus, in general pattern; each has a prom-
inent process on the seventh tergite, and a process is also present
on the eighth tergite of N. recta Banks. The female sternites are
apparently unmodified in all four species. Names proposed by Banks
have been applied to this material, but until Banks' types can be
critically studied there will be some uncertainty as to the correctness
of these assignments.

Neoperla atripennis Banks. Figure 20.

Neoperla (Javanita) atripennis Banks, 1924, Bull. Mus. Comp. Zool., 65, (12),
p. 428, pi. 2, fig. 21.

JEWETT: STONEFLIES FROM THE PHILIPPINES 83

The dark wings and body and bicolored legs distinguish this spe-
cies. Banks' illustration of the male genitalia is fairly accurate,
though what appears to be a plate on the eighth tergite is only a
dark sclerotized area. The head bears rather small eyes and ocelli
(fig. 20).

Fig. 19. Leuctra (Rhopalopsole) pala-
wana, female genitalia of holotype, ven-
tral view.

Fig. 20. Neoperla atripennis, female
head and prothorax.

Banks erred in placing this species in the subgenus Javanita. The
"last ventral segment" of the male is not "pointed in the middle,"
though Banks may have been led to this conclusion by distortion
of his specimen in drying. This species is related to the other Philip-
pine members of the genus. The male forewing measures about 11
mm. in length, that of the female 16 mm.

Material examined. — Mindanao: Mount McKinley (east slope;
3,000 ft. alt.), Davao Province; 1 9 ; Aug. 29, 1946; F. G. Werner.

Surigao; 2 cf (USNM); C. F. Baker.

Neoperla obliqua Banks. Figure 21.

Neoperla obliqua Banks, 1913, Proc. Ent. Soc. Washington, 15: 172, pi. 8,
fig. 12.

The oblique crossvein between the radial sector and the radius
seems always to be present in the forewing (occasionally distinct
only in one wing) . The apical portion of the wings is without cross-
veins. Figure 21 illustrates the head pattern. The male genitalia
do not include a process on the eighth tergite but are otherwise sim-
ilar to those of N. recta Banks. The male forewing measures 14-17
mm. in length, that of the female 15-23 mm.

Material examined. — Mindanao: Mount McKinley (east slope;
3,000-5,200 ft. alt.); 17 d\ 10 9 (7 d\ 3 9, SGJ); in mossy and

84

FIELDIANA: ZOOLOGY, VOLUME 42

open forests and on rocks in stream; Aug. 21-Oct. 20, 1946; H. Hoog-
straal and F. G. Werner.

Mount Apo (east slope); alt. 2,800 ft. (Todaya), 4,300 ft. (Camp
Mainit), 6,000 ft. (Camp Meran); 5 d" , 54 9 (1 o", 15 9, SGJ);
Nov., 1946; H. Hoogstraal and G. Alcasid.

Fig. 21. Neoperla obliqua, male head
and prothorax.

Fig. 22. Neoperla oculata, female
head and prothorax.

Galog River, Mount Apo (6,000 ft. alt.) ; 1 9 (USNM, labeled
"Paratype N. apoana"); Oct. 18; C. F. Baker.

Burungkot, Upi, Cotabato Province (1,500 ft. alt.); 2 9 (1 9 ,
SGJ), at light; Dec. 31, 1946- Jan. 1, 1947; F. G. Werner.

Tangcolan, Bukidnon Province; 1 9 (USNM); C. F. Baker.

Agusan, Bukidnon Province (1,000 ft. alt.); 1 9, July 26, 1946;
H. Hoogstraal and D. Heyneman.

Luzon: Mount Makiling; 1 &, 2 9 (USNM); C. F. Baker.

Neoperla oculata Banks. Figure 22.

Neoperla oculata Banks, 1924, Bull. Mus. Comp. Zool., 65: 428, pi. 2, fig. 16.

The shape of the head, especially the visible suture lines behind
the ocelli, and the very large eyes separate this species (fig. 22). The
male genitalia are very similar to those of N. recta. As in N. obliqua
the crossvein between the radial sector and radius is slanted obliquely
inward. The third segment of the maxillary palpus is noticeably
curved inward, differing in this respect from the other species of
Neoperla treated in this paper. This is also the smallest of the spe-
cies studied. The length of the forewing of the male is about 11 mm.,
that of the female 12-14 mm.

Material examined. — Mindanao: Badiang Tagabuli, Santa Cruz,
Davao Province (2,000 ft. alt.); 16 9 (6 9 , SGJ); at light; Dec. 10,
1946; M. Celestino.

JEWETT: STONEFLIES FROM THE PHILIPPINES 85

Maco, Tagum, Davao Province (near sea level) ; 7 9 (2 9 , SGJ) ;
at light, in original forest (3 9 , at Sitio Taglawig, near Maco);
Oct. 7-15, 1946; H. Hoogstraal.

Surigao; 2 d" (USNM); C. F. Baker.

Neoperla recta Banks

Neoperla recta Banks, 1913, Proc. Ent. Soc. Washington, 15: 172, pi. 8, fig. 10;
1937, Philipp. Jour. Sci., 63: 135.

This species is readily distinguished by the straight crossvein
between the radial sector and radius and by the crossveins in the
apical part of the forewings. In the male there is also a prominent
process on the eighth tergite as well as one on the seventh tergite.
The eyes are somewhat more prominent than those of N. obliqua
Banks. The shape and morphology of the rear median area are very
similar to those of N. obliqua. The length of the male forewing is
13-20 mm., that of the female 16 mm. (from Luzon only). Speci-
mens from Luzon are much smaller than those from Mindanao.

Material examined. — Mindanao : Camp Meran, Mount Apo (east
slope; 6,000 ft. alt.); 4 d1 (2 d\ SGJ); "rocks in swift stream;"
Nov. 6, 1946; H. Hoogstraal.

Luzon: Los Bafios; 1 & (USNM); C. F. Baker.

Mount Makiling; 3^,1$ (USNM); C. F. Baker.

Imugin, northern Viscaya; 1 9 (USNM); C. F. Baker.

Genus Phanoperla Banks

Phanoperla Banks, 1938, Jour. Fed. Malay States Mus., 18: 221.

There are at least two species of this genus in the Philippines.
They are separable on the basis of the male genitalia and to some
extent on the distance between the ocelli. The wing venation of the
two cannot be differentiated. In both, the origin of Cu2 is usually
slightly ahead of the first cubito-medial crossvein, but sometimes it
is at or slightly behind it. The rear median portion of the head
appears to be identical in the two species.

Phanoperla bakeri (Banks). Figure 23.

Neoperla bakeri Banks, 1924, Bull. Mus. Comp. Zool., 65: 426-427, pi. 2,
fig. 19.

Figure 23 shows the male genitalia of what is considered to be
P. bakeri. Banks' illustration, cited above, is closer to what I con-
sider to be his N. Clarissa. Specimens examined match neither his
written description nor his illustration; possibly the legends for his

86 FIELDIANA: ZOOLOGY, VOLUME 42

figures 15 (N. consimilis Banks) and 19 were transposed. In speci-
mens identified as P. bakeri the anterior point on the male genital
hook is always rounded, and there is one large depressed patch of
spicules on the ninth tergite. In the male the ocelli are rather widely
separated. The male forewing measures 10-14 mm., that of the

Fig. 23. Phanoperla bakeri, male Fig. 24. Phanoperla clarissa: A, male

genitalia, dorsal view. head and prothorax; B, male genitalia,

dorsal view.

female 16 mm. The males from Luzon are smaller than those from
Mindanao.

Material examined. — Mindanao: Mount McKinley (east slope;
6,400 ft. alt.) ; 1 d1 ; in mossy forest; Sept. 7, 1946; H. Hoogstraal.

Mount Apo (east slope; alt. 4,300 ft., Camp Mainit; 6,000 ft.,
Camp Meran) ; 1 d\ 2 9 (1 d\ 1 9, SGJ) ; on rocks in swift stream,
and at light; Nov., 1946; H. Hoogstraal.

Luzon: Mount Makiling; 3 d* (USNM); C. F. Baker.

Phanoperla clarissa (Banks). Figure 24.

Ochthopetina clarissa Banks, 1913, Trans. Amer. Ent. Soc, 39: 204.

The ocelli seem to be consistently closer together in this species,
especially in the male, than in P. bakeri, and the anterior tips of the
male genital hooks are pointed (fig. 24) . Spicules on the ninth tergite
of the male are arranged in two elevated patches, not in a single
depressed patch as in P. bakeri. This species is of the same general
size as P. bakeri, and material from Mindanao is somewhat larger
than that from Luzon.

Material examined. — Palawan: Bacungan, near Puerto Princesa
(near sea level); 1 c? (SGJ); "near small rocky stream;" Mar. 26,
1947; F. G. Werner.

Busuanga (Calamianes Island group): Dimaniang (near sea
level); 1 cf; "on shrub leaf over creek;" Mar. 14, 1947; H. Hoog-
straal.

JEWETT: STONEFLIES FROM THE PHILIPPINES 87

Mindanao: Burungkot, Upi, Cotabato Province (alt. 1,500 ft.);
1 <? (SGJ); at light; Jan. 1, 1947; F. G. Werner.

Sitio Taglawig, near Maco, Tagum (near sea level); 1 <?; "at
light in forest;" Oct. 11, 1946; H. Hoogstraal.

Dansalan, Lanao; 1 d\ 1 9 ; May 16, 1946; H. Hoogstraal.

Kolombugan; 1 d1 (USNM); C. F. Baker.

Surigao; 1 c? (USNM); C. F. Baker.

Luzon: Mount Makiling; 2 d" , 1 9 (USNM); C. F. Baker.

REFERENCES

Banks, Nathan

1913. On a collection of neuropteroid insects from the Philippine Islands.
Proc. Ent. Soc. Washington, 15: 170-180, pis. 8-9 (Perlidae, pp. 171-172).

1920. New neuropteroid insects. Bull. Mus. Comp. Zool., 64: 297-362, 7 pis.
(Perlidae, pp. 314-325).

1924. Descriptions of new neuropteroid insects. Bull. Mus. Comp. Zool., 65:
419-455, 4 pis. (Perlidae, pp. 426-428).

1937. Philippine neuropteroid insects. Philipp. Jour. Sci., 63: 125-174, 6 pis.
(Perlidae, pp. 134-137).

1938. New Malayan neuropteroid insects. Jour. Fed. Malay States Mus., 18:
220-237, 17 figs. (Perlidae, pp. 221-223).

Hanson, John F.

1941. Studies on the Plecoptera of North America, II. Bull. Brooklyn Ent.
Soc, 38: 57-66.

Hoogstraal, Harry

1951. Philippine Zoological Expedition, 1946-1947. Narrative and itinerary.
Fieldiana: Zool., 33: 1-86, 7 figs., 7 pis.

Kimmins, D. E.

1950. Some new species of Asiatic Plecoptera. Ann. Mag. Nat. Hist., (12),
3: 177-192.

Klapalek, Fr.

1912. Sauter's Formosa-Ausbeute. Plecoptera. Ent. Mitt., 1: 342-351.

1921. PIScopteres nouveaux. 3rd pt. Ann. Soc. Ent. Belg., 61: 320-327.

Navas, (Rev. Pere) Longino

1918. Insecta nova. Mem. Pont. Accad. Romana Nuovi Lincei, (2), 4: 1-22
(Perlidae, pp. 3, 4).

Needham, James G.

1905. New genera and species of Perlidae. Proc. Biol. Soc. Washington, 18:
107-110.

PlCTET, FRANQOIS JULES

1853-57. TraitS de Paleontologie. 2nd ed., 4 vols, and atlas. Paris.

Ricker, W. E.

1952. Systematic studies in Plecoptera. Ind. Univ. Publ., Sci. Ser., no. 18,
pp. 1-200.

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