Zoologica Scripta. Vol. 19, N o . 1 , pp. 73-99, 1990 030(!-3256/90 $3.00+ .

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01990 The Norwcgian Academy of Science and L e t t e r s

A contribution to the Dicyrtomidae (Collembola) of Hawaii

RICHARD J. SNIDER
Act eprerl 2 1 November I989

Snider. K.J . 1990. A contribution t o the Dicyrtomidae (Collembola) of Hawaii.-Zool. Scr. 19:
73-99.
Ten new species in three subgenera of Dicyrtonia arc described from the Hawaiian Islands.
Specimens were received from collcctions made on Hawaii. Maui, Kauai and Oahu. Species
definitions are based on chactotaxy of head, legs and the circumanal region. In addition, presence
o r absence of latcral spines (neosminthurid sctae) on the parafurcular lobes may assist in grouping
species within subgenera. Previous records of dicyrtomids from Hawaii include only one species,
Ptenorlirix (Pupirioidos) duhia (Folsom). The following new species are described: Ptenorhrix
(Ptenothrix) hawaiicneis sp.n., Pfenorlirix (Papirioides) kauuiensis sp.n., P. (Pupirioides) serruiu
sp.n., Ilicyrtomu (C’ulvutorninu) sylvestrurilis sp.n., L). (Culvutotninu) hrevifihru sp.n., D. (C‘ulva-
tornir~cc)tesssseluru sp.n., D. (Culvu/ornina) longidigiiu sp.n., D. (Calvaiomina)bellingeri sp.n., D.
(Culvutomitiu) made.stris sp.n.. and D. (Culvutoniinu) microdentutu sp.n.
H..I. Snider, Uepartmeni o] Zoologv, Michipti Stare Univesiiy, Eust Lansing, MI 48824, U.S.A .

Introduction manubrium (parafurcular lobes) appears to be a valuable

taxonomic feature. This characteristic and the lack of
The purpose of this report is to describe 10 new species of vertical cephalic spines permitted him to place Dicyrto-
dicyrtomids from Hawaii. Specimens were accumulated mina oryzae Murphy, 1960 in the “rufescens group”.
from field samples and research holdings either collected Huther (1065, 1967) found the same characteristic for
or borrowed by Drs K. A. Christiansen and P. F. Bell- Dicyrtominu rufescens (Reuter) and Dicyrtomina pseudo-
inger as the basis of their forthcoming monograph on the rufescens Huther. Yosii & Lee (1963) devised a scheme
collembolan fauna of Hawaii. The species disclosed here for denoting the setae of the anogenital region of Ptenoth-
were sent to the author for examination and description. rix, which Yosii (1969) later modified for use with Dicyv-
The Hawaiian Collembola have not received much tomina and Calvutomina. Yosii & Ashraf (1965) used
attention. although thc island group has attracted large these characteristics and other setae of the head and body
numbers of biologists. Carpenter (1004) was the first to to clearly separate Dicyrtoma (Dicyrtoma) pakistanica
publish on the collembolan fauna of the Hawaiian islands Yosii & Ashraf from Ptenothrix and Dicyrtomina.
(Sandwich Islands). H e described five species, none of Folsom (1924) described a new species and genus,
which were dicyrtomids. Folsom (1932) made the next Papirioides jacobsoni from Sumatra. Its general mor-
contribution to the faunal list by adding 22 new records. phology was close to Ptenothrix, except it exhibited a
He described Ptenothrix duhia from a small collection large abdominal, median dorsal, club-like structure. This
(four specimens) considering it to be similar to, or “may characteristic seemed sufficiently unique for generic
be Ptenothrix mirahilis Denis (1929) . . . described from recognition. Uchida (1957b) placed his species Pupir-
China”. Because Denis had only four specimens, of which ioides uenoi in this genus for the same reasons. Stach
two lacked the dorsal tubercle, the early status of P. (1965), agreeing with this characteristic, described Papir-
duhia, in Folsom’s opinion (1932), remained in doubt. ioides aequituherculata from North Korea. He com-
The most recent work is Zimmerman’s (1948) “Insects of mented that Folsom’s (1932) dubia and Denis’ (1929)
Hawaii”; a rewrite of Folsom (1932), presented without mirahilis belonged in Papirioides.
species descriptions and following the higher categories Uchida (1944) described Dicyrtomina trukana from
presented by Wormersley (1939). Micronesia, a species with a dorsal abdominal tubercle
Modern work on the Dicyrtomidae began with Stach and 5 segmented antenna. He also compared three genera
(1957). He recognized the need for comparative chaeto- of sminthurids exhibiting tubercles with Dicyrtomina.
taxy studies to separate similar species. In addition he Stach (1957) placed trukana in the subgenus Pseudo-
redefined the family, organized a key to genera and dicyrtomina based on the 5 segmented antenna, however,
species, and presented a number of critical descriptions of the genus otherwise appeared to fit well within Calvato-
European examples. Following his publication, a number mina. Betsch (1980) followed Yosii (1969) by placing
of significant contributions were made by authors in the Dicyrtomina verrucosa Handschin, Dicyrtomina hom-
Pacific region. Uchida (1953, 1957~)proposed a dental hayensis Yosii and D. trukana in the genus Calvatomina
setae formula for dicyrtomid species. Subsequently, a under the “Pseudodicyrtomina group”. He noted that
number of modifications of his system have come into use. only trukana appeared to exhibit 5 antenna1 segments. All
Murphy (1960) stated that presence or absence of three species showed a posterior abdominal tubercle with
lateral spines (neosminthurid setae) at the base of the strong spines. Yosii (1969) separated Calvatomina,
73 Zoologica Scriptu 19
74 R . J . Snider

Table I . Ncw species from Huwuii described in this pup‘’‘ Table 11, C‘ircirmunul untipur(rJirrcu1ur .rerujhrmiilue wlirre ( + ) = hlurit
spine. ( - ) = wta-like. (i-) = sharp &pineund ( 0 ) = .setu uhsent; 312 = 3
Family Dicyrtomidae spines. 2 . s ~ t u - l i kS~f 0. = 5 spines und Of5 = spines uh.sen/
~~~~ . -... . ~ _ _
Tribe Ptcnothricini Species Anogcnital sctac I’arafurcular
setae
Prenorhrix (f‘tmothri.r) huwuiiensis, sp.n. ... .
Ptenothrix (Pupirioide.5) kauaiensis. sp.n.
M M ’ N N’ A 0 T H 3
‘ L
Ptetiothrix (I’upirioidrs) serrutu, sp.n.
hu wuiirwsis + + + O + + + + + 015
Tribe Dieyrtornini
huuaiensis + + + o - + + - - 015
serrutu + + + o + - - c - + 015
“formosanu group” sy 1vestru tilis + + + + - - + - + 510
Dicyr/omlr (C’uivutominu)sylvestru/ili.s, sp.n. hrevijihru + + + + - o + + + 312
Dicyrromu (Culvutrmiinu) brevifihru, sp.n. tesselutu + + + - - ( I + + + 312
Dicyrtomu (C‘ulvutomiriu) tesst~lutu.sp.n. longidigitu + + + * - - + - + 312
Dicyrtomu ((‘ulvutominu)lorigiciigitu, sp. n . hrlliri,y~ri + + + + - - + * + 312
Uicyrtotnu (Chlvutominu)hellingeri. sp.n. mudr~srris + - + + + - + O + 510
“riifi‘secris group” microden/u/cr + - + - + - + - + 510
Dicyrtomu ((‘ulvurorninu) mudestris, sp.n .
Dicyr/onru (Culi~utorninu)microdentutu, sp.n

“Pseudodicyrtominm group” from the “rufescens group” taken with their interpretation. Presence or absence of
and “forrnosana group” on the basis of the ventral dental bothriothrix A and D is a strong characteristic and along
setae; 4-2-1-1-1 . . . 1 for the former and 4-2-1-1 . . . 1 with the number and type of spines on the metatibia can
for the latter two groups. Betsch (1980) said that the be used to clearly separate Ptenothricini from Dicyrto-
“Pseudodicyrtomina group” would seem to be equivalent mini. In addition, at the generic or subgcneric level, the
to Pupirioides with respect to Ptenothrix. So far, the following characteristics used by them for North Ameri-
collections from Hawaii have not produced any speci- can species cannot bc applied successfully to Pacific
mens from the “Pseudodicyrtomina group”. species: antennal subannulation, number of unpaired
Currently we rely upon three modern works to position facial setae [extreme variation between Dicyrtornu.
higher categories within Dicyrtomidac. First, Richards Dicyrtornina. Calvutornina and Ptenothrix, (Snider,
(1968) separated Bothriovulsus, Papiriodes and Ptenoth- 198Su,h)], or claw structure.
rix into the tribe Ptenothricini on the basis of bothriothrix Even though a strong argument can be presented for
D being present and antennal segments 111 and IV being separation of subgenera into genera a s presented by
subannulated. He placed Dicyrtomu and Dicyrfomina in Betsch (1980), a more conservative system incorporating
the tribe Dieyrtomini based on lack of bothriothrix D and his ideas and Christiansen & Bellinger (1981) and Yosii
subannulations on antennal segment IV. His opinions (1969) is used here. Ten new species are described from
were predicated on thcsis research concluded 12 years Hawaii. Their separation from each other and from
earlier (Richards, 1956). Very little editing between the- species described earlier from the Pacific-tropical area
sis publication and 1968 had taken place. It is, therefore, relies heavily on anogenital chaetotaxy.
not surprising that the papers by Yosii (lY6O,lY66,1969), Table I lists the new species and, where appropriate,
Yosii & Ashraf (1965) and Yosii & Lee (1963) were not the “group” to which they belong. Types are deposited in
included in his final analysis. While he cited Stach (1957) the Bishop Museum, Honolulu, Hawaii. In addition,
for elevating Dicyrtomidae to family level, he did not Table I1 summarizes the anogenital chaetotaxy used for
mention t h e genera Pseudodicyrtornina (Stach, 1957) or species division.
Gihherathrix (Uchida, 1 9 5 2 ~ )In. spite of these omissions,
his analysis o f genera was strongly presented. Many
critical morphological characteristics commonly used Account of species
today are a direct result of his research.
Betsch (1980) presented an overview of the Dicyrtomi- Ptenothrix (Ptenothrix) hawaiiensis sp.n. (Figs 1-31)
dae. He combined the work of Richards (1968), Yosii
(1960, 1966, 1969), Yosii & Lee (1963) and Yosii & Material exumined. Holotype ( 9 )and paratype on didcs. Hawaii,
Hawaii County, Lava Tree State Park, beating vegetation, 20 January
Ashraf (1965) to produce analytical reviews of each genus IYX2, K. A. Christiansen. Maui, Maui County, Koolau Gap, Halcakala
and an updated version of Richards’ (1968) taxonomic Crater. 7000 feet clcv., moss on rock. 7 February 1964, Tsuda.
key to genera. Betsch believed that Calvatornina and
other subgenera should be elevated to generic status. H e Description
pointed out that Calvatorninu formed a homogeneous
Holotropical group, while Dicyrtornina was Holarctic in Color. (Description based on specimens mounted on
distribution. slides in Hoyer’s medium.) Background opaque to white
Christiansen & Bellinger (1981) did not include the with blue pigment distributed in the following patterns.
work o f Betsch (1980) in their treatment of the North Antenna uniformly blue. Vertex of head with blue macu-
American collembolan fauna. As a result they took a lae, dark blue-black patch between ocelli; frons and gena
conservative view of the family. They preferred to use a with irregular blue mosaics. Great abdomen with blue
hybridization of Richards (1968) and Gisin (1960) which mosaics forming irregular lateral patterns. Legs with blue
reduced Bothriovulsus, Dicyrtornina and Ptenorhrix to banding at intervals. Furcula light blue. Lesser abdomen
subgenera under Dicyrtomu. Some exception must be blue with flecks of blue-black (Fig. 1).
Zoologicu Scriptu 19
 Dicyrtornidne (Collembola) of’Hawaii 75

serrate, E setae ratio EI/E2 = 0.754.95 and E3/E2 =

1.80-1.85 (Fig. 27). Mucro with inner and outer teeth
(Fig. 28). Parafurcular lobes with normal, smooth setae
(Fig. 29). Circumanal setae M, M’ and N spine-like and
smooth, seta sa normal; other setae follow pattern
M N T H G A0 sa
3 2 + + + + -
other upper valve setae finely serrate; female subanal
appendage slightly curving, acuminate (Fig. 30). Both-
riothrix D present. Anterior body setae (AA-EE) spine-
like, smooth (Fig. 31), posterior dorsal setae short,
dagger-like becoming longer laterally. Length up to
2 mm.
Fig. I . I’/e~io/hris(P/rno/hri,r)huwuiiensis sp.n., hahitus. Total length
up t o 2 m m .
Diagnosis. Ptenothrix (Ptenothrix) hawaiiensis sp.n. most
closely resembles Ptenothrix (Ptenothrix) ciliophora
+
Morphology. Eyes 8 8; ocellus H 1/2 diameter of others (Yosii & Lee, 1963) and Ptenothrix (Ptenothrix) saxutilis
(Fig. 2). Mean antenna1 ratio 1 :4.2:5.5: 1 ; ANT I with 5 (Yosii & Lee, 1963), both endemic to Korea. Besides
dorsal and 2 ventral setae (Fig. 3); ANT I1 with 2 dorsal having facial setae 1-1-2-2-1-1, their labral setal formula
and 1 ventral cup sensilla; ANT 111 divided into 8 distal is unique: 6/5, 3, 4. However, ciliophora has all median
subsegments, armed with cup sensilla, subapical sensilla facial setae of the same diameter, while saxatilis shares
exposed, lying in shallow depressions; ANT IV divided the same thick type seta 1-1 followed by 2-2-1-1 as found
into 4 subsegments. Outer maxillary lobe consists of in huwaiiensis. They all share finely serrate setae on the
simple palp and 1 sublobal hair (Fig. 4). Labrum with upper anal valve. Color pattern will easily separate these
setal pattern 6/5,3,4 (Fig. 5 ) . Dorsal setae of head spine- species. Another species, Ptenothrix (Ptenothrix)gracilli-
like (Fig. 6): 2 unpaired facial setae (Fig. 7) and l + l oval cornis (Schaffer) has 1-1-2-2-1-1 facial setae. Further, if
organs on lower frons. the illustration given by Yosii (1969) is re-examined, his
Foreleg. Coxa, trochanter and femur not clearly seen; formula 6/3, 5 , 4 may be interpreted as 61.5, 3, 4 which
tibiotarsus with 4 cup sensilla and I oval organ on anterior would place this species close to hawaiiensis. The species
surface (Fig. 8), 1 oval organ on posterior surface (Fig. 9); can be separated on the basis of anal N setae, grucillicor-
pretarsus with anterior and posterior setulae; unguis lacks nis exhibits a setaceous rather than blunt spine-like type.
tunica, with 2 strong inner teeth; unguiculus with strong It is clear that huwaiiensis can be classified between the
corner tooth, apical filament reaching beyond tip of “southern” and “Holoarctic” groups of Ptenothrix sugges-
unguis, knobbed (Fig. 10). ted by Yosii (1969). He utilized 1-1 blunt facial setae and
Mcsoleg. Coxa with 3 anterior setae and 1 “courte relative lengths of circumanal setae. His system allows
epine” (Fig. 11); trochanter with 4 anterior and 1 pos- comparison with Ptenothrix (Ptenothrix) brasiliensis
terior setae and oval organ (Fig. 12); femur with 8 (Delamare & Massoud), 1963 and Ptenothrix (Ptenothrix)
anterior and 5 posterior setac, 1 posterior setula and oval argentina (Delamare & Massoud), 1963 based o n the
organ (Fig. 13); tibiotarsus with 4 cup sensilla and 1 oval length of sa and sa’; huwaiiensis is comparable. Cephalic,
organ on anterior surface (Fig. 14), 1 “courte epine” on tibia1 and labral setae patterns will separate the species.
posterior surface (Fig. 15); pretarsus with anterior and Within the Hawaiian species complex, superficial com-
posterior setulae; unguis lacks tunica. with 2 strong inner parison to Ptenothrix (Pupirioides) kauaiensis sp.n. will
teeth; unguiculus with strong corner tooth, and some- distinguish it from that species by color pattern, facial
times a second accessory tooth, apical filament knobbed, spines, labral setae, lack of hump, and circumanal setal
reaching beyond tip of unguiculus (Fig. 16). pattern.
Metaleg. Coxa with 4 anterior setae and 1 “courte
epine” (Fig. 17); trochanter with 5 anterior and 1 pos-
terior sctae (Fig. 18); femur with 9 anterior and 3 pos-
terior setae and oval organ (Fig. 19); tibiotarsus with 5 Ptenothrix (Pupirioides) kuuaiensis sp.n. (Figs 3 2 4 5 )
cup sensilla and 1 “courte epine” on anterior surface (Fig.
Mu/eria/ exurnitzed. Holotype ( 9 ) and 5 paratypes on slides. Collec-
20), 1 “courte Cpine” and 2 differentiated setae o n pos- tion data: Hawaii. Kauai County, Wailua River crossing. swccping ncar
terior surface (Fig. 21); pretarsus with anterior and pos- pavcd road t o Experiment Station. I8 January 1067, P. F. Bellingcr.
terior setulae; unguis lacks tunica, with 2 strong inner
teeth; unguiculus with strong corner tooth, and some- Description
times with second accessory tooth, apical filament
knobbed, rcaching beyond tip of unguis (Fig. 22). Color. The specimens examined are all on slides; their
Great abdomen. Collophore with 1 1 subapical and+ color pattern is poorly preserved. However, Dr Christian-
I + 1 lateral setae (Fig. 23), sacs warty. Corpus of tenacu- sen made notes on the pattern before mounting at least
lum with 4 setulae, ramus with 3 teeth and basal horn (Fig. one example. Yellow with purple pigment limited to
+
24). Manubrium with 7 7 dorsal setae (Fig. 25). Dens middorsal line on posterior portion of great abdomen,
with 3. 2, I , . . . 1 Ve setae (Fig. 26), dorsal setae basally with small scattered patches elsewhere o n head and body.
7oologicu Scrrptu 19
76 R . J . Snider

13

14 15

19
\

li 10 20 21
Figs 2-22. Ptenothrix (Ptenothrix) hawaiiensis sp.n.-2. Left eyepatch.-.?. Antenna1 segment 1 . d . Outcr maxillary lobe.--5. L a b r u m . 4 . Dorsal
setae of head.-7. Median facial setae.-8. Foretibiotarsus. anterior surface.--9. Foretibiotarsus, posterior surface.--10. Foreclaw.-/l.
Mcsocoxa.-/2. Mesotrochantcr.--I3. Mesofeniur.--I4. Mesotibiotarsus. anterior surface.-fS. Mesotibiotarsus. posterior surface.--16.
Mesoclaw.-/7. Metacoxa.--IS. Metatrochiinter.-lY. Metafemur.-20. Metatibiotarsus, anterior surface.-2-1. Metatibiotarsus, posterior
surface .-22. Metaclaw .

Zoologicu Scriptu 19
 Dicyrtomidae (Cullembola) of Hawaii 77

Figs 2.ZG.11. Ptenothrix (Plenorhrix) huwuiiensis sp.n.-23. Collophorc, latcral vicw.-24. Corpus of retinaculurn.-25. Mucro.-26. Dens, ventral
surface.-27. Dens, dorsal surfacc and detail.-211. Manubrium, dorsal surface.-29. Parafurcular setac.--30. Anal papilla.-31. Anterior body
s c t x and bothriotrix A .

Antenna uniformly purple. Legs banded with purple Mesoleg. Coxa with 3 anterior setae and 1 “courte
(Fig. 32). 6pine” (Fig. 44); trochanter with 4 anterior and 1 pos-
terior setae, D2 seta much reduced (Fig. 45); femur with
+
Morphology. Eyes 8 8; ocelli D and H subequal and 112 anterior and posterior “courtes epines” , cup sensillum on
diameter of others (Fig. 33). Mean antenna1 ratio outer margin (Fig. 46); tibiotarsus with 5 cup sensilla and
4: 19:24:6; ANT I with 5 dorsal and 2 ventral setae (Fig. 3 “courtes epines” on anterior surface (Fig. 47), no
34); ANT TI with 2 dorsal and 1 ventral cup sensilla; ANT
111 divided into 7 subsegments, armed with cup sensilla,
subapical sensilla exposed, lying in shallow depressions;
ANT IV divided into 5 subsegments. Outer maxillary
lobe consists of simple palp and sublobal hair (Fig. 35).
Labrum with setal pattern 6/7,2,4 (Fig. 36). Dorsal setae
of head short, spine-like, a-seta reduced; facial setae not
truly blunt, arranged as 1-1-2-2-1-1, lower frons with
A”
1 + 1 oval organ (Fig. 37).
Foreleg. Coxa with 1 seta, no oval organ (Fig. 38);
trochanter 2 anterior and 2 posterior setae (Fig. 39);
femur with basal posterior and distal anterior oval organs,
cup sensillum on outer margin (Fig. 40); tibiotarsus with 4
cup sensilla and 3 oval organs (almost “courtes @pines”)
on anterior surface (Fig. 41), oval organ lacking on
posterior surface (Fig. 42); pretarsus with anterior and
posterior setulae; unguis lacks tunica, with basal lateral
teeth, 2 inner teeth, well developed; unguiculus with
32
corner tooth, subapical filament reaching beyond tip of Fig. 32. Prenorhrix (Pupirioidrs) kauuiensis sp.n., habitus. Total length
unguis, knobbed (Fig. 43). of entire animal up to 1.5 m m .

Zoologica S c r i p 19
78 R. J . Snider

' i

33 34

Fig.s f'tmorhriu (f'upirioides) kuuuiensis sp.n.-.3.3. Left eycpatch.-.34. Antcnnal segment l.-.35. Outer maxillary lobe.-.36. I,abrum.-.?7.
Median facial sctiic.-.W. Forecoxa.-.W. Foretrochantcr.--40. F o r c f c m u r . 4 1 . Foretibiotarsus, anterior surfacc.-42. Forctibiotarsus, posterior
surfacc.4.1. F o r c c l a w . 4 4 . Mcsocoxa.4-5. Mcsotrochantcr.46. M c s o f c m u r . 4 7 . Mcsotibiotarsus, antcrior s u r f a c c . 4 8 . Mcsotibiotarsus,
posterior surfacc.--.lY. Mcsoclaw.,SO. Mctacoxa-SI. Mctatrochantcr.-S2. Mctafcrnur.-SS. Mctatibiotnrsus, anterior surfacc.-S4. Mctatibio-
tarsus. posterior surface.--5.5. Metaclaw.

Zoologicu Scriptu 19
 Dicyrtomidae (Collembola) of Hawaii 79

61
Fix.s 5 6 4 5 . l’/cm/hrix (Papirioidrs) krriruicri.si.s sp.n.-S6. Collophorc, lateral view.-S7. Corpus of rctinacu1um.-SK. Manubrium, dorsal
surfacc.-FY. Dens. ventral s u r f a c e . 4 0 . Dens. dorsal surface and d c t a i l . 4 l . M u c r o . 4 2 . Anal papilla, f c r n a l e . 4 3 . Female subanal
appendage . 4 3 . Dorsal chactotaxy of a h d o m c n . d . 5 . Parafurcular sctac

“courtes epines” on posterior surface (Fig. 48); pretarsus Female subanal appendage acuminate, curved (Fig. 63).
with anterior and posterior setulae; unguis similar to Bothriothrix D present. Trunk setae AA and CC small,
foreclaw, unguiculus with subapical filament reaching setaceous; BB, EE, GG, HH spine-like. Dorsal projec-
well beyond tip of unguis, knobbed (Fig. 49). tion not well defined (Fig. 64). Parafurcular setae seti-
Metaleg. Coxa with 4 anterior setae and 1 “courte form, except 1 spine-like anterior example (Fig. 65).
Cpine” (Fig. SO); trochanter with 5 anterior and 1 pos- Length up to 1.5 mm.
terior setae, D2 seta reduced (Fig. 51); femur with an-
terior and posterior “courtes Cpines” and outer cup sen- Diagnosis. Ptenothrix (Papirioides) kauaiensis sp .n .
silla (Fig. 52); tibiotarsus with 5 cup sensilla and 3 appears to stand alone in the subgenus by having the
“courtes epines” on anterior surface (Fig. 53), posterior unguicular subapical filament knobbed on all legs.
surface with 2 heavily serrate differentiated setae (Fig. Further, the female subanal appendage is acuminate and
54); pretarsus with anterior and posterior setulae; unguis curving; other species have short blunt setae. Among the
with basal lateral teeth, 2 inner teeth; unguiculus with Hawaiian representatives of the genus, the spine-like
strong corner tooth, subapical filament reaching tip of trunk macrochaetae, unguicular filaments and subanal
unguiculus, knobbed (Fig. 55). appendages will separate kauaiensis from all others. Until
Great abdomen. Collophore with 1 + 1 subapical and more material becomes available, placement in Papir-
+
1 1 lateral setae (Fig. 56), sacs warty. Corpus of tenacu- ioides sp. must remain in doubt. The low, poorly defined
lum with 4 setulae, ramus with 3 teeth and basal horn (Fig. hump and strong Ptenothrix sp. characteristics seem to
57). Manubrium with 8 + 8 dorsal setae (Fig. 58). Dens position this species nearer that subgenus.
with 3-2-1 . . . 1 Ve setae (Fig. 59), dorsal E and L setae
serrate (Fig. 60), EI/E2= 0.7M.90 and E2/E3= 2.00-
2.80. Mucro with inner and outer teeth, outer file irregu- Ptenothrix (Pupirioides)serrutu sp.n. (Figs 66-100)
lar and conical, inner even and rounded (Fig. 61). Circu-
manal setae M, M’, and N spine-like and smooth, seta sa Marcrial uxamirzed. Holotypc ( 9 ) and 5 paratypes o n slide; Hawaii,
Honolulu County, Oahu. Mt. Tantalus, tO00 fcct clcv., pan trap,
normal; other setae follow pattern (Fig. 62) Novcmbcr 1966, J . Vockeroth. Additional paratypcs (all o n slides);
Maui County, Maui, Halcakala Crater. Koalua Gap, 7000 feet elev.,
M N T H G A. sa moss o n rock, 7 Fcbruary 1964, Tsuda, collector, 6 specimens o n slide I ,
3 2 - t - t - - - 2 spccimcns o n slidc 2; Maui, lao Valley State Park, approximately I

Zoolofiicu Scriptu 19
80 R. .I.
Snider

milc from “needle”. scruti forest and fcrns and grass, under bark o l cup sensilla on outer margin (Fig. 81); tibiotarsus with 5
fallen trees and stones. 3 February 1982, K. A . Christianscn, I spcci- cup sensilla on outer margin and 3 “courtes Cpines” on
mcn; Honolulu County. Oehu, Wnimca Arborcturn. IS0 feet clcv.,
. ” domcsticatcd plant section, on dccorativc lava rock in pools,
H,nw‘iiian anterior surface (Fig. 82), posterior surface with 1 cup
27 Fcbruary 19x2. K . A . Christianscn. sensillum and no “courte Cpine” (Fig. 83); pretarsus with
inner and outer setulae; unguis lacking tunica, with basal
Llescription lateral teeth and 2 well-developed inner teeth; unguiculus
with corner tooth, subapical filament reaching tip of
Color. (Description based on specimens mounted on unguis (Fig. 84).
slides in Hoyer’s medium.) Background yellow; antenna Metaleg. Coxa with 4 anterior setae and 1 “courte
purple-blue except distal and basal quarters of ANT TI, epine” (Fig. 8s); trochanter with 5 anterior and 1 pos-
which are light; band of purple on lower frons, with terior setae, D2 normal (Fig. 86); femur with basal pos-
irregular mosaics concentrated on gena; macula of blue terior oval organ and anterior distal “courte Cpine” and
between eyepatches, otherwise light. Body irregularly outer cup sensilla (Fig. 87); tibiotarsus with 5 cup sensilla
mottled, with faint wash of blue, middorsal line on an- and 3 “courtes epines” on anterior surface (Fig. 88),
terior half, lower anal valves bluish, upper valve distally posterior surface with 2 heavily serrate differential setae
blue. Legs banded. Manubrium-dens purple-blue, be- (Fig. 89); pretarsus with anterior and posterior setulae;
coming darker distally; mucro blue (Fig. 66). unguis with basal lateral teeth, 2 outer teeth, and 2 well-
developed inner teeth; unguiculus with corner tooth,
Morphology. Eyes 8 + 8; ocellus D smaller than others lamella slightly more expanded than on foreclaw or
(Fig. 67). Mean antenna ratio 1 :4.5:5: 1; ANT I with 5 mesoclaw, subapical filament reaching tip of unguis (Fig.
dorsal and 2 ventral setae (Fig. 68); ANT I1 with 1 dorsal 90).
and 2 ventral cup sensilla; ANT Ill divided into 7 distal Great abdomen. Collophore with 1 + 1 subapical and 1
subsegments, armed with cup sensilla, subapical sensilla + 1 lateral sctae (Fig. 91), sacs warty. Corpus of tenacu-
exposed, lying in shallow depressiohs; ANT IV divided lum with 5 4 setulae, ramus with 3 teeth and basal horn
into 5 subsegments. Outer maxillary lobe consists of +
(Fig. 92). Manubrium with 8 8 dorsal setae (Fig. 93).
simple palp and 1 sublobal hair (Fig. 69). Labrum with Dens with 3 , 2, 1 , 1 . . . 1 Ve setae, dorsal E and L setae
setal pattern 6/7,2,4 (Fig. 70). Dorsal setae of head short, serrate (Fig. 94), EI/E2 = l.S(k2.35 and ED/E2= 2.60-
spine-like (Fig. 71); 5-7 unpaired facial setae (Fig. 72) 3.40 (Fig. 95). Mucro with numerous fine inner and outer
+
1 + 1 “courtes epines” and 1 1 oval organs on lower teeth (Fig. 96). Circumanal setae M, M’ and N spine-like
frons. and smooth, seta sa normal; other setae follow pattern
Foreleg. Coxa with 1 anterior seta (Fig. 73); trochanter
M N T H G A(, sa
with 2 anterior and 2 posterior setae (Fig. 74); femur with
basal posterior and distal anterior oval organs (anterior
3 2 - + - + -
qualifies as “courte Cpine”), cup sensillum on outer mar- (Fig. 97). Female subanal appendage straight, short,
gin (Fig. 7.5); tibiotarsus with 4 cup sensilla and 1 “courte blunt (Fig. 98). Bothriothrix D present. Trunk setae,
6pine” on anterior surface (Fig. 76), oval organ lacking large, strong, only DD spine-like. Dorsal projection well
on posterior surface (Fig. 77); pretarsus with anterior and defined, low (Fig. 99). Parafurcular setae setiform (Fig.
posterior setulae; unguis lacks tunica, with basal lateral 100). Length up to 2.2 mm.
teeth, 2 inner teeth; unguiculus with corner tooth, subapi-
cal filament just reaching beyond tip of unguis (Fig. 78). Diagnosis. Ptenothrix (Papirioides) serratu sp.n. unlike
Mesoleg. Coxa with 3 anterior setae and 1 “courte kauaiensis, lacks knobbed subapical filaments on the
epine” (Fig. 79); trochanter with 4 anterior and 1 pos- unguiculi and has a greater number of unpaired facial
terior setae, D2 seta normal (Fig. 80); femur with basal setae. Ptenothrix (Papirioides) duhia (Folsom, 1932) is
posterior oval organ and distal anterior “courte Cpine”, easily separated from serrata on the basis of color pattern,
anal setae and prominent hump. Further, duhiu has a 1-
1-1-1-2-1-1 facial setal formula, while serrata has 1-1-1-
2-1-1-1. Another difference is found in the setae of ANT
I; dubiu has 4 spine-like dorsal setae and 2 posterior,
serrata has 5 dorsal, normal setae. The only other species
that superficially resembles serrata is Ptenothrix (Pupir-
ioides) mirabilis (Denis, 1929) from China. That species
has individuals that exhibit both the presence and absence
of dorsal humps. Furthermore, the dorsal setae of the
dens lack serrations in mirahilis.

Dicyrtoma (Calvatomina)madestris sp.n. (Figs 101-134)

Material examined. Holotypc ( P ) and 37 paratypes in ethanol: Haw-

aii, Kauai county, Kauai, gladc by Wailua River, abovc falls, damp
grass, I6 January 1967, 5 additional paratypes on slide from same
Fig. 66. Pfenorhrix (Papirioides)serrata sp.n., habitus. Total lcngth up location, 4 specimens on slide and 18 in ethanol collected from sticks on
to 2.2. mm. ground; Alakai Swamp Trail, forcst moss on logs, 17 January 1967;

Zoologica S c r i p I9
 Dicyrtomidue (Collembolu) of Hawuii 81

67 % 71

4
72
/

F3 79

84

J
76 77 90 82 83 89
Figs 67-90. P&nothri,t (Pupirioides) serrufu s p . n . 4 7 . Right e y e p a t c h . 4 8 . Antcnnal segment I . d Y . Outer maxillary lobe.-70. Labrum.-7I.
Dorsal sctac of head.-72. Median facial sctac.-73. Forecoxa.-74. Foretrochanter.-7.5. Forefemur.-76. Foretibiotarsus, anterior surface.-77.
Forctibiotarsus, posterior surface.-78. Foreclaw.-7Y. Mesocoxa.4U. Mcsotrochanter.41. M c s o f e m u r . 4 2 . Mcsotibiotarsus, anterior
surfacc.4.7. Mcsotibiotarsus, posterior s u r f a c c . 4 4 . M e s o c l a w . 4 5 . M c t a c o x a . 4 6 . Mctatrochanter.47. M c t a f e m u r . 4 8 . Mctatibiotarsus,
anterior s u r f a c c . 4 9 . Mctatibiotarsus, posterior surface.--90. Metaclaw.

Zoologicu Scriptu I9
 R. J . Snider

I I
1 I

i
I
I
1 I
1
I
I
I
I
I
I
I

92
1
I

ill
93
95

*-

98

\ 97 100
F ~ g sYl-100. I’tcrrothrix (Puprrioidcs) .serrutu sp.n.-Y/. Collophorc.-Y2. Corpus of rctinaculum.-9.?’. Manuhrium. dorsal surface.-94. Dens.-
9.7. Dcntal sctac E,-E2.-96. Mucro.--Y7. Anal papilla. fernalc.-W Fcmalc subanal appendage.-YY. Chactotaxy of abdomen, lateral vicw.-/OO.
Parafurcular sctae.

Wailua River. crossing on paved road. inland from Experiment Station, lying in shallow depressions; ANT IV not subsegmented.
rotten wood, soil and vegetation. 18 January 1967, 4 spccimcns o n slide
and 5 in ethanol: Mauna Kapu, Waianac Mountains, litter under Dorsal setae of head small, not spine-like; 5 unpaired
Cu.siturinusp.. 20 October 1966, P. F. Bellinger. median facial setae and oval organs lacking (Fig. 105);
outcr maxillary lobe consists of simple palp and 1 sublobal
Description hair (Fig. 106). Labrum with setal pattern 6/5, 5 , 4 (Fig.
107).
Color. Background cream to light yellow with purple Foreleg. Coxa without scta or oval organ (Fig. 108);
pigment dusting and irregular polygons. Head with intcr- trochanter with 3 anterior and 1 posterior setae (Fig. 109);
ocular macula of light purple; light polygons o n frons with femur with posterior oval organ and distal anterior
macula between antennal bases and slightly below; purple “courte @pine”,and outer cup sensilla (Fig. 110); tibiotar-
increases on gena and becomes heaviest on post-gena; sus with 4 cup scnsilla and 1 “courte epine” on anterior
antenna with ANT I light purple, ANT IT becoming dark
distally, ANT 111 and IV dark purple. Body with thoracic
segmentation outlined in light purplc; dorsally with a faint
colorless stripe originating at the first thoracic segment
and terminating 3/4 distance of abdomen; posterior ab-
dominal region light purple with setal sockets colorless;
laterally thorax to abdomen with dark purple, inter-
sperscd with irregular maculac; lesser abdomen dusted
with light purplc; legs light purplc; furcula cream with
light purplc dusting at base of dens (Figs 101, 102).

+
Morphology. Eyes 8 8; ocellus D 1/2 diameter of E, all
other ocelli cubequal (Fig. 103). Mean antennal ratio
4 : 13: 18:4; ANT 1 with 4 dorsal and 3 ventral setae (Fig. 102 ‘%&2
104); ANT 11 with 2 dorsal and 1 ventral cup sensilla; Figs 101-102. Dicyrtornu (Culvutomina) vzudtWris sp.n.-101. Habitus,
ANT I11 not subsegmented, subapical sensilla exposed, latcral view.-IO2. Habitus, dorsal view. Total length up t o 1 mm.

Zoolo~rcrr >Scrrptu19
 Dicyrtomidue (Collembola) of Huwuii 83

b 7 p 10.7-129. Dicyrfoma (Calvumnina)mudrstris sp.n.--lO.?. Left eycpatch.-l04. Antenna1 segment I.-1M. Facial setac.--lM. Outer maxillary
lobc.--107. Labrum.--IOX. Forccoxa.-109. Forctrochantcr.-ll0. Forcfemur.--ill. Forctibiotarsus, anterior surfacc.-l12. Forctibiotarsus,
posterior surfacc.-l13. Foreclaw.--ill. Mesocoxa.--llS. Mcsotrochantcr.-116. Mcsofemur.--117. Mcsotibiotarsus, anterior surfacc.-!IR.
Mcsotibiotarsus, posterior surface.-l 19. Mcsoclaw.-l20. Mctacoxa.-lZ/. Mctatrochanter.-122. Metafemur.--I2.?. Mctatibiotarsus, antcrior
surfacc.-124. Mctatibiotarsus. posterior surface.--125. Metaclaw.--I26. Collophore.-l27. Manubrium, dorsal surface.-128. Dens, ventral
surface.-129. Dens, dorsal surface.

Zoologica Scriptu I9
84 R. J . Snider

Dicyrtoma (Calvatomina) sylvestratilis SQ.~.

(Figs 135-1 67)

Mu/cviul c~.rtirrrinctl.Holotypc ( 5’) and 7 paratypcs m slides. Hawaii.

Kauai County, Kauai. Wailua-Hanalei Trail, 2 milcs from pavcd road on
Wailua side. rotten wood. fern litter and under bark, 18 January 1967.
.
dratypc;
13. bamc location. swecping, 3 paratypcs. P. F. Bcllingcr; Hono-
lulu County, Oahu, Mi. Tantalus, 1000 feet clcv.. pan trap, Acuciu sp.
and (’ercus sp., 22 Octoher 1966, J . Vockcroth.

Description

surface (Fig. I 1 I ) , posterior surface with 1 “courte epine” Color. (Description based on specimens mounted on
(Fig. 112); prctarsus with anterior and posterior setula; slides in Hoyer’s medium.) Background yellow to opaque
unguis with 1 outer tooth, tunica and pseudonychia; with purple distributed in the following patterns. Head
unguiculus with well-developed corner tooth, subapical with rose-purple reaching below eyepatches, dark macula
filament 1/2 length of inner margin of unguis, reaching between eyes, remaining area yellow-cream; antenna
beyond its tip (Fig. 113). rose-purple basally becoming darker distally. Body with
Mesoleg. Coxa with 2 anterior and 1 posterior setae purple-brown reaching laterally, sometimes dorsum very
(Fig. 114); trochantcr with 4 anterior and 1 posterior setae light, giving the appearance of lateral stripes; lesser abdo-
(Fig. 115); femur with posterior oval organ and distal men cream; legs with light dusting of rose-purple (Fig.
“courte kpine” and outer cup sensillurn (Fig. 116); tibio- 135).
tarsus with 4 cup sensilla and 3 “courtes Cpines” on
anterior surface (Fig. 117), posterior surface with 1 Morphology. Eyes 8 + 8, ocelli subequal (Fig. 136).
“courte epine” (Fig. 11 8); pretarsus with anterior and Mean antcnnal ratio 4: 20 :23 :5 ; ANT I with 5 dorsal and
posterior setulae; unguis with outer tooth, 1 inner tooth, 2 ventral setae (Fig. 137); ANT I1 with 2 dorsal and I
tunica and pseudonychia; unguiculus with corner tooth, ventral cup sensilla; ANT I11 not subsegmented, armed
subapical filament short, 1/5 length of inner margin of with cup sensilla, subapical sensilla exposed, lying in
unguis (Fig. 119). shallow depressions; ANT IV not divided into sub-
Metaleg. Coxa with 3 anterior setae and 1 “courte segments. Outer maxillary lobe consists of simple palp
epine” (Fig. 120); trochanter with 4 anterior and 1 pos- and 1 sublobal hair (Fig. 138). Labrum with setal pattern
terior setac (Fig. 121); femur with posterior oval organ 6/S, 5 , 4. Dorsal setae of head reduced; 5-7 unpaired
and distal “courte epine”, outer cup sensillum (Fig. 122); facial setae (Fig. 139).
tibiotarsus with 5 cup sensilla and 3 “courtes kpines” on Foreleg. Coxa with 1 seta (Fig. 141); trochanter with 2
anterior surface (Fig. 123) posterior surface with 1 anterior and 2 posterior setulae (Fig. 141); femur with 5
“courte Cpine” (Fig. 124); pretarsus with anterior and anterior and’s posterior setae, 2 posterior setulae, outer
posterior setulae; unguis with outer tooth, 1 inner tooth, cup sensillurn, posterior basal oval organ elongated, an-
tunica and pseudonychia; unguiculus with corner tooth terior distal “courte @pine”(Fig. 142); tibiotarsus with 4
(sometimes 2) and occasionally a distal tooth (sometimes cup sensilla and 3 “courtes Cpines” on anterior surface
2j, subapical filament 1/4 length of inner margin of unguis (Fig. 143), 1 “courte Cpine” on posterior surface (Fig.
(Fig. 125). 144); prctarsus with anterior and posterior setulae; unguis
+
Great abdomen. Collophore with 1 1 subapical and with pseudonychia, tunica, outer tooth, 2 small inner
1 + 1 lateral setae (Fig. 126), sacs warty. Corpus of teeth; unguiculus with small corner tooth, subapical fila-
tenaculum with 4 setulae, ramus with 3 teeth and basal ment tapering, about length of lamella, reaching beyond
horn. Manubrium with 10 + 10 dorsal setae (Fig. 127). tip of unguis (Fig. 145).
Dens with 4,2, I , I . , I Ve setae (Fig. IZS), dorsal E and
,

L setae all smooth (Fig. 129). E,/E2= 1.0-1.25, E3/E2 =

0.00-1.10. Mucro with numerous inner and outer teeth,
irregular (Fig. 130). Circumanal setae which follows pat-
tern for subgenus M’ is reduced and finely setaceous
M N A. T H G and L
+ + + - + o +
(Figs 131, 132). Female subanal appendage acuminate,
slightly curved (Fig. 133). Parafucular setae commonly
with 5 ( 6 ) spine-like ciliated setae (Fig. 134). Length up to
1 mm.

Diugtiosrs. Dicyrtomu (Culvatominu) madestris sp.n . I\ a a

member of the “rufescens group” of Yo\ii, 1969. It comes
closest to Dicyrtomu (Calvutominu) yueyarnensiJ Yosii, 135
?’he differencesbetween the two species are ‘Isted F I ~11’5 Ilrc)r/orncr (CulLaroncmu) ~ j h c ~ c t r u r r lsp
r ~ n , habitu\, lateral
in Table 111. view T o t d length up t o I I mm

Loolopcii .k rcptu 19
 Dicyrtomidue (Collembolu) Of’ Hawaii 85

T;iblc IV. Morplio/ogicu/ dij~cvenrcvberween Dicyrtoma (Calvatomina) sylvcstratilis .sp.tt. . Dicyrtoma (C:alvatoniina) solomonensis (Yosii ), und
Dicyrtonis (Calvatomina) modcsta ( Yosii)
.sJlveSri.urili.s .solomoncn.sis rYIode.sru
~ - __ ~ ~ ~ .~ __ ~~ ~ ~~ ~~~ - ~~ ~ ~

Macrochactac M. N . N ’ blunt, of N ‘ 112 length of” and blunt N ’ 112 length of N and sctaccous N’112 length 01 N and almost
equal length setacenus
G a c t a sctaccous Ycs Yes N o , blunt
A ? scta smallcr than A4 Suhcqunl Yes Y cs
Dorsal dcntal sctac 71.. IOE, D, rcduccd ( D , 11.1 10I2, IOE. D , normal ( D I and D: IOL. OE, D, normal (D, and Dz
lc11gth of D2) su bcq uil I ) su bcq ual)

Mesoleg. Coxa with 3 setae and 1 ”courte Cpine” (Fig. Dicyrtoma (Calvatomina) brev$bra sp-n. (Figs 168-192)
146); trochanter with 3 anterior and 1 posterior sctae (Fig.
Ma/eriul exuminerf. Holotype ( 9 ) and 2 paratypc on slidcs. Hawaii,
147); femur with 7 anterior setae and 2 posterior setulae, Kalawao County, Molokai, vicinity of Kalahuapuco lookout, scrub
outer cup sensillum, posterior basal oval organ elongated, forcst, 12 January 19x3, K . A . Christianscn. holotypc; Hawaii County,
anterior distal “courte Cpine” (Fig. 148); tibiotarsus with Hawaii. Kukuihaclc, unidentified bird’s nest. 14 Dcccnihcr 1961. Wil-
son, collector, paratypc. Honolulu County, Oahu, Manoa Falls trail,
5 cup sensilla and 3 “courtes epines” on anterior surface flats near beginning. litter. 23 September 1966, P. F. Bcllingcr.
(Fig. 149), 1 “courte Cpine” on posterior surface (Fig.
150); pretarsus with anterior and posterior setulae; unguis Doscription
with pseudonychia, tunica, outer tooth, 2 inner teeth;
unguiculus with corner tooth, subapical filament short, Color. (Description based on specimen preserved in etha-
reaching tip of unguiculus (Fig. 151). nol.) Background white with dark blue pattern. ANT 1
Metaleg. Coxa with 3 setae and 1 “courte Cpine” (Fig. with distal blue ring; ANT 11 light blue, becoming dark
152); trochanter with 4 anterior and 1 posterior sctae (Fig. blue distally; ANT I11 light blue basal 1/4, dark blue
153); femur with 8 anterior and 2 posterior setac, 1 distally; ANT IV dark blue. Vertex of head light blue with
posterior setula, posterior oval organ elongated, distal interocular macula dark blue, extending just below anten-
anterior “courtc epine” and outer “courte Cpine” (Fig. nal bases; upper frons with light blue macula, lower frons
154); tibiotarsus with 5 cup sensilla and 3 “courtes Cpines” to labrum blue, surrounding area white; gena with dark
on anterior surface (Fig. ISS), posterior surface with I bluc forming a stripe that extends to occiput. with a stripe
“courte epine” (Fig. 156); pretarsus with anterior and of blue extending from antennal bases to mouthparts.
posterior setulae: unguis with pseudonychia, tunica. Thoracic segments laterally blue, forming horizontal
outer tooth, 2 inner teeth; unguiculus, with corner tooth, stripes dorsally, segments lined with white and scattered
subapical filament short, reaching beyond tip of unguis white maculae. Abdomen dark bluc with dark blue stripe
(Fig. 157). originating at thorax and terminating at apex before anal
Great abdomen. Collophore with 1 + 1 subapical and valves, this stripe is bordered with white; laterally with
1 + 1 lateral setae, sacs warty. Corpus o f tenaculum with scattered white maculae roughly forming a square; anal
4 setae, ramus with 3 teeth and basal horn (Fig. 158). valves dorsally light with dark apical macula. Manubrium
+
Manubrium with 9 9 dorsal setae (Fig. 159). Dens with light blue, dens white. Collophore light blue. Legs with
4,2,1. 1 . . . 1 Ve setae (Fig. 160), dorsal E and L setae all coxa and trochanter dark blue, femur dark blue distally,
smooth (Fig. 161), EI/E2= 1.20-1.40, E3/E1 = 1.30 (Fig. otherwise white, tibiotarsus light blue distal 3/4 (Figs
162). Mucro with numerous inner and outer teeth (Fig. 168-1 70).
163). Circumanal setae follow pattern for subgenus
Morphology. Eyes8 + 8, ocelli subequal (Fig. 171). Mean
M N A,) T H G and L M’
antennal ratio 4 : 15:21 :5; ANT I with 5 dorsal and 2
+ + - - + - + + ventral setae (Fig. 172); ANT I1 with 2 dorsal and 1
(Fig. 164); female subanal appendage straight, slightly ventral cup sensilla; ANT TI1 not subsegmented, armed
curved and sharply tapered at apex. Parafurcular setae with cup sensilla, subapical sensilla exposed. lying in
spine-like and ciliated (Fig. 165). Bothriothrix A and C shallow depressions; ANT TV not divided into subseg-
very reduced, B normal (Fig. 166). Body setae fine in ments. Dorsal setae of head reduced; 5 unpaired facial
thoracic region, becoming spine-like posteriorly (Fig. setae (Fig. 173).
167). Length up to 1 . 1 mm. Foreleg. (Only claw adequate for description.) Pretar-
sus with anterior and posterior setulae; unguis with pseu-
Diagnosis. Dicyrtorna (Culvutomina) sylvestratilis sp.n. in donychia, tunica, 1 outer tooth and single inner tooth;
many respects resembles Dicyrtoma (Culvutomina) solo- unguiculus without corner tooth, subapical filament
rnotietzsis (Yosii, 1969) and Dicyrtoma (Calvatomirzu) tapering beyond tip of unguis (Fig. 174).
modrsfu (Yosii, l969), both from the Solomon Islands. Mesoleg. Coxa with 3 anterior setae; femur with 5
Clearly they share the same labral setal pattern, 4 tentacu- anterior and 5 posterior setae, with outer cup sensillurn,
lar sctae, 1 + 1 and 1 + 1 collophore setae, Ve setal anterior and posterior setulae (Fig. 175); (tibiotarsus
pattern 4 , 2 , 1 , 1 . . . 1 and inner margin of unguis with 1 , inadequate); pretarsus with anterior and posterior setu-
2, 2, teeth. However the circumanal macrochaetae pat- lac; unguis with pseudonychia, tunica, 2 outer teeth and 2
tern niay be used to separate the 3 species as shown in fine inner teeth; unguiculus with corner tooth, subapical
Table IV. filament tapering beyond tip of unguis (Fig. 176).
Zoologicu Scriptu 19
86 K. J . Snider
 Dicyrtomidue (Collemholu) o j Hawaii 87

192). Bothriothrix A and C normal, B missing, rep-

resented by cup sensillum only, D missing. Body setae i n
thoracic region tine, becomine spine-like posteriorly.
Length up to 1 mm.

Iliagnosis. Dicyrtorna (Calvatominu) brevifihru sp. n. is

allied to the forrnosanu group of Yosii (1969) by virtue of
the M, M‘, N , N‘ setal pattern, as well as similar parafur-
cular setal pattern. However, the presence of dorsal cup
+
sensilla ( 1 1 ) on the manubrium, 7L, 9 E dorsal setae o n
dens, and striking color pattern will separate the species
from the f o r m o m n u group and Hawaiian species. Further
collecting will n o doubt improve the dcscription pre-
sented here.

Dicyrtoma (Calvatomina) tesselata sp.n. (Figs 193-225)

Matcriul eutrrriiric4. Holotypc ( P ) and 5 paratypes o n slides. Hawaii,

Honolulu County. Oahu, below I’ali above Anouai PI.. upper Manoa
Vallcy, dead lcavcs among loose rocks, 30 Novcmber 1966, P. F.
Bcllingcr (holotypc). Oahu (30 Novcmber 1966). 2 o n slide. Oahu. Mt.
170 T;tntalus, rotten wood in bamboo stand. 16 Scptemhcr 1966, P. F.
Bcllingcr, 1 (8). O a h u , Manoa Falls trail. flats near beginning. litter, 23
Fig.s 168-170. Dicyrrornu ((blvutonrir~u)hrei.ifihru s p . n - / 6 K . Hobitus. September 1966, P. F. Bellingcr, 2 on slide.
lateral vicw.-/69. Hahitus. d o r d vicw.-/70. Hcad. frontal view.
Description

Metaleg. Coxa with 4 anterior setae (Fig. 177); tro- Color. (Description based on specimens mounted on
chanter with 4 anterior and 1 posterior setae (Fig. 178); slides in Hoyer’s medium.) Background yellow with pur-
femur with 8 anterior and 3 posterior setae, basal oval ple pigment laid down in irregular polygons. Antenna
organ, distal setula and outer cup sensillum (Fig. 179); purple. Vertex of head with dark purple macula between
tibiotarsus with 5 cup sensilla and 3 “courtes @pines”on eyepatches, light purple stripe connected to dark macula
anterior surface (Fig. lXO), posterior surface with 1 of frontal interocular area; lateral purple band on gena
“courte dpine” (Fig. 181); pretarsus with antcrior and with light suffusion of purple. Body with dark purple
posterior setulae; unguis with pseudonychia, tunica, 2 lateral bands that join near the postero-dorsal area, anal
outer teeth and 2 inner tine teeth; unguiculus with corner papilla light purple. Legs with light purple, distal portion
tooth, subapical filament tapering, short and fine, reach- of femur darker. Furcula light purple on manubrium and
ing beyond tip of unguis (Fig. 182). dens, mucro with small amounts of purple (Fig. 193).
Great abdomen. Collophore with 1 + 1 subapical and
+
1 1 lateral setae, sacs warty (Fig. 183). Corpus of Morphology. Eyes 8 8, ocelli approximately subequal+
tenaculum with 4 setae, ramus with 3 teeth and basal horn (Fig. 194). Mean antenna1 ratio 4: 18.5:24:6.5; ANT I
(Fig. 184). Manubrium with 9 + 9 dorsal setae and 1 + 1 with 5 dorsal and 2 ventral setae (Fig. 195); ANT I1 with 2
cup sensilla (Fig. 185). Dens with 4 , 2 , 1, 1 . . . 1 Ve setae
dorsal and 1 ventral cup sensilla; ANT 111 not subseg-
(Fig. 186), dorsal E and L setae smooth (Fig. 187), EI/E2 mented, armed with cup sensilla, subapical sensilla ex-
= 0.81, E3/E2 = 1.60 (Fig. 188). Mucro with numerous posed, lying in shallow depressions; ANT IV not subseg-
inner and outer teeth (Fig. 189). Circumanal setae follow mented. Outer maxillary lobe consists of simple palp and
the pattern for the genus 1 sublobal hair (Fig. 196). Labrum with setal pattern 6 / 5 ,
5 , 4. Dorsal setae of head reduced; 5-6 unpaired facial
M N A,, ‘T H G and L M’ setae (Fig. 197).
+ + - - + + + + Foreleg. Coxa with 1 anterior seta (Fig. 198); tro-
chanter with 2 anterior and 1 posterior setae (Fig. 199);
(Fig. 190). N’ short, spine-like; female subanal append- femur with 5 anterior and 5 posterior setae, outer cup
age acuminate, slightly curved (Fig. 191). Parafureular sensillum, posterior setula and basal oval organ (Fig.
area with 3 spine-like, ciliated setae and 2 setaceous (Fig. 200); tibiotarsus with 5 cup sensilla and 3 “courtes epines”

Figs 130-134. Ilicyrtomu (C’ulvtrrominu) rnadesrris sp.n-/30. Mucro.-/31. Anal papilla, dorsal view.-1.<2. Anal papilla, latcrnl vicw.-1.3.3.
Fcinalc subanal appendagc.-/34. Parafurcular sctac.
Figs 136-167. Dicyrtornu ( C h l v a ~ o m i n nsy/vc,sirutilis
) sp.n.-/.?6. Left cycpatch.-137. Antcnnal scgmcnt I .-/SK. Outer maxillary lobe.-/.W.
Facial scta.-/40. Forccoxa.--14/. Foretrochantcr.-/42. Forcfemur.-/43. Forctibiotarsus, anterior surfacc.-/44. Foretibiotarsus, posterior
surface.-/4.5. Foreclaw.--146. Mesocoxa.-/47. Mesotrochanter.-/48. Mcsofcmur.--I4Y. Mcsotibiotarsus. anterior surfacc.--150. Mesotihio-
tarsus, posterior surface.-/5/. Mcsoclaw.--ISZ. Mctacoxa.-/S.?. Metatrochantcr.-154. Mctafemur.-/55. Mctatihiotarsus, anterior surfkc.-
1.56. Mctatihiotarsus. posterior surfacc.-l57. Metaclaw.-ZW. Corpus of rctinaculum.--15Y. Manubrium. dorsal surfacc.-I60. Dens, ventral
surfacc.-/6/. Dens, dorsal surfacc.-Z62. Dental sctac EI-E3.--163. Mucro.-/64. Anal papilla, fcmalc.-/65. Parafurcular sctac.-166.
Bothriotrix A-B.--167. Body sctae, thoracic region.

Zoologica Scriptu 19
88 R . J . Snider

172
177

175

Q i
/ ?/ '0, I70

174

181 180

/ y' 179

I
I
I I
I I
184

185

186 187 188 192

Figs /7/-/92. Dicyrroma (Calvaromina) brevifihra sp.n.-/71. Left eyepatch.-/72. Antenna1 segment 1.-/73. Unpaired facial s c t a e . 4 7 4 .
Forcclaw.--17S. Mesofcmur.-/76. Mcsoclaw.-177. Metacoxa.--178. Metatrochantcr.--I7Y. Mctafcmur.--l8U. Mctatibiotarsus, anterior
surfacc.-/8/. Metatibiotarsus, posterior surface.--I82. Mctaclaw.-/83. Collophore.-/84. Corpus of retinaculum.-I85. Manubrium, dorsal
surfacc.-/M. Dens. ventral surface.--187. Dens, dorsal surface.-/88. Dental sctae EI-E3.-/8Y. Mucro.-IYO. Anal papilla, male.--191. Female
subanal appendage.-/Y2. Parafurcular setae.

Zoologica Scripru 19
 Dicyrtornidae (Collemhola) o f Hawaii 89

Diagnosis. Dicyrtomcc (Calvatomina) tesselatcz sp.n.

strongly resembles Dicyrtorna (Cnlvatomina) hrevifihra
sp.n. in many respects. It also has many characteristics of
the ‘:formosnna group”. Separation from the Hawaiian
members of the genus is by 6L, 9E setae and D, < Dz on
the dens; in addition. hrevifihra lacks a corner tooth on
the fore-unguiculus and has 1 + 1 cup sensilla o n the
manubrium. The strong color pattern alone will dis-
tinguish this \pecies from other Hawaiian dicyrtomids.

Dicyrtoma (Calvatomina)microdentata sp.n.

(Figs 226-237)
.
big 19.3 Ihc‘yrlomu ((‘ulvutoniiriu) t c w ~ l u r a\p n hahitti\. lateral VICW.
Mulerrul c ~ ~ u n i r r i rHolotypc
d ( 0 ) and paratypc on one didc Hdwaii,
Totdl length up to I mm
Kaudi County, Kauai, Kokce Stdte Park, swept from long grass by road.
17 January 1967, P F. Bcllingcr

on anterior surface (Fig. 201), posterior surface with 1 Description

“courte epine” (Fig. 202); pretarsus with anterior and
posterior setulae; unguis with pseudonychia, tunica, Color. (Description based on specimen mounted on slide
sometimes second outer tooth and 1 inner tooth; unguicu- in Hoyer’s medium.) Background cream-yellow with light
lus with corner tooth, subapical filament about 1/2 length purple pigment. Antenna light purple, becoming darker
of inner edge of unguis, tapering just beyond tip of unguis distally. Head with light dusting of purple on vertex, gena
(Fig. 203). and postgenal areas. Body with fine granules of purple
Mesoleg. Coxa with 3 anterior setae and 1 “courte dorsally, becoming darker laterally, giving the appear-
tpine” (Fig. 204); trochanter with 3 anterior and 2 pos- ance of lateral bands. Legs light purple. Furcula cream
terior setae (Fig. 205); femur with 10 anterior and 2 (Fig. 226).
posterior setae, outer cup sensillum, and basal oval organ
(Fig. 206); tibiotarsus with 5 cup sensilla and 3 “courtes Morphology. Eyes 8 + 8, ocelli A and B subequal, slightly
epines” on anterior surface (Fig. 207), 1 “courte ipine” larger than others (Fig. 227). Mean antennal ratio
on posterior surface (Fig. 208); pretarsus with anterior 4:19:21:6. ANT I with 5 dorsal and 2 ventral setae;
and posterior setulae; unguis with pseudonychia, tunica, antennal segments 111 and IV unsubsegmented, other
outer tooth and 2 small inner teeth; unguiculus with features not clear. Dorsal setae of head fine; 5 unpaired
corner tooth, subapical filament short, fine, reaching just facial setac. Outer maxillary lobe and labrum not seen.
beyond tip of unguis (Fig. 209). Coxae, trochanters, femora, and tibiotarsi not clear
Metaleg. Coxa with 4 anterior setae and 1 “courte enough for illustration.
Cpine” (Fig. 210); trochanter with 4 anterior and 1 small, Foreleg. Tibiotarsus with 4 cup sensilla, 3 “courtes
fine posterior setae (Fig. 21 1); femur with 10 anterior and epines” on anterior surface, 1 “courte epine” on pos-
1 posterior setae, outer cup sensillurn, 2 posterior setulae terior; pretarsus with anterior and posterior setulae;
and basal oval organ (Fig. 212); tibiotarsus with 5 cup unguis with pseudonychia, tunica, 1 outer and 2 inner
sensilla, 3 “courtes Cpines” on anterior surface (Fig. 213), teeth; unguiculus with corner tooth and subapical fila-
1 “courte Cpine” on posterior surface (Fig. 214); pretarsus ment over 1/2 length of inner edge of unguis, reaching
with anterior and posterior setulae; unguis with pseudo- beyond its tip (Fig. 228).
nychia, tunica, sometimes outer tooth, subapical filament Mesoleg and Metaleg.Tibiotarsus with 5 cup sensilla, 3
short, fine, tapering just beyond tip of unguis (Fig. 215). “courtes epines” o n anterior surface, posterior with 1
Great abdomen. Collophore with 1 1 subapical and 1+ “courte Cpine” ; pretarsus with anterior and posterior
+ 1 lateral setae, sacs warty (Fig. 216). Corpus of tenacu- setulae; unguis with pseudonychia, tunica, 1 outer and 2
lum with 4 setae, ramus with 3 teeth and basal horn (Fig. inner teeth; unguiculus with corner tooth, subapical fila-
217). Manubrium with 9 + 9setae (Fig. 218). Dens with4, ment short, fine, about 1/3 length of inner margin of
2, 1 , 1 . . . 1 Ve setae (Fig. 219), dorsal E and L setae all unguis (Fig. 229).
smooth (Fig. 220), EI/E2= 0.95-1.10, E3/E2= 1.10-1.30 Great abdomen. Collophore with 1 1 subapical and +
(Fig. 221). Mucro with numerous inner and outer teeth +
1 1 lateral setae (Fig. 230), sacs warty. Corpus of
(Fig. 222). Circumanal setae follow the pattern tenaculum with 4 setulae, ramus with 3 teeth and basal
M N A. T H G and L M’ horn (Fig. 231). Manubrium with 9 + 9 dorsal setae (Fig.
+ + - - + + + + 232). Dens with 4, 2, 1, 1 . . . 1 Ve setae, dorsal setae
smooth, 8L, YE, E setae ratio EI/EZ= 1.10 and E3/E2 =
(Fig. 223); female subanal appendage heavy, sharp, 1.20 (Fig. 223). Mucro with numerous inner and outer
tapered and curved at apex (Fig. 224). Parafurcular setal teeth, teeth of inner margin much smaller than outer (Fig.
pattern 3 spine-like and 2 setaceous (Fig. 225). Bothrioth- 234). Circumanal setae follow pattern
rix A , B, C normal, D missing. Body setae fine in thoracic
region, becoming spine-like posteriorly. Length up to M N A,) T H G and L M’
1 mm. + + + - + - + -
Zoologico Scripto 19
90 R . J . Snider

(Fig. 235); female subanal appendage acuminate, curving Table V. Morphological d i j & w z c e s heiween Dicyrtoma (Calvatomina)
(Fig. 236). Parafucular setal pattern 5 spine-like, ciliated longidigita s p . n . , Dicyrtoma (Calvatomina) brevifibra .sp.n., Dicyrtoma
(Calvatomina) sylvestratilis sp.n. and Dicyrtoma (Calvatomina)
setae (Fig. 237). Bothriothrix A , B, C normal, D missing. tesselat a sp .n .
Body setae fine in thoracic region, becoming spine-like
posteriorly. Length up to 1 mm. lorigidigita hreviJihra sylvestrurilis resselata
. .... .. ~~ ~

Circumanal T scta Yes Absent Yes Absent

Diagnosis. Dicyrtoma (Calvatominu) microdentata sp. n. present and
setaceous
like Dicyrtoma (Calvatomina) madestris sp.n. falls within Parafurcular setec 3 3 5 3
the “rufescens group” of Yosii (1969). The species may be spine-like
separated on the basis of ungual configuration of the Subapical filament to 1 :1 112 1:3 1 z2.75
outer edgc of
metalegs, manubrial setae, number of E, L setae on dens, met aunguicul us
and circumanal setae G. Another member of the
“group”, Dicyrtoma (Calvatominu)pugoda (Yosii, 1966)
exhibits most of the characteristics of microdentata. It can Cpine” on posterior surface (Fig. 250); pretarsus with
be separated by having 10L, 10E dental setae, lacking G anterior and posterior setulae; unguis with pseudonychia,
seta on the anal valve and a distinctive color pattern. tunica, outer tooth and 2 inner teeth; unguiculus with
Dicyrtoma (Calvatomina) yaeyamensis (Yosii, 1965) is strong corner tooth and small second tooth, subapical
very similar, but can be distinguished from microdentata filament heavy, tapering, as long as unguiculus (Fig. 251).
on the basis of color pattern, setaceous M’ seta, blunt T +
Great abdomen. Collophore with 1 1 subapical and 1
seta and uniform teeth on mucro. The unique G seta, + 1 lateral setae, sacs warty. Corpus of tenaculum with 4
reduced to a setulae makes this species easy to identify. setulae, ramus with 3 teeth and basal horn. Manubrium
not clear. Dens with 4 , 2 , I , 1 . . . 1 Ve setae, other setae
obscure, E and L setae smooth, EI/E2 = 1.00, E3/E2 =
1.10. Mucro with numerous inner and outer teeth (Fig.
Dicyrtoma (Calvatomina)longidigita sp.n. (Figs 238-256) 252). Circumanal setae follow pattern
Material exuminrd. Holotypc (9) on slide. Hawaii, Hawaii County, M N A,, T H G and M’ N’ L
Hawaii, Puu Pilc Kohala Mountains. clev. 4500 feet, forest floor debris,
h July 1964. Haaa.
+ + - - + - + + +
(Fig. 253); female subanal appendage heavy, curved and
Description sharply tapered at apex (Fig. 254). Parafucular setae 3
spine-like and 2 setaceous (Fig. 255). Bothriothrix D
Color. Purple with irregular light maculae. Appendages missing. Body setae fine in thoracic region, becoming
dark purple (Fig. 238). spine-like posteriorly (Fig. 256). Length up to 2 mm.

Morphology. Eyes 8 + 8, subequal (Fig. 239). ANT I with Diagnosis. Dicyrtoma (Calvatomina)longidigita sp.n. is a
5 dorsal and 2 ventral setae (Fig. 240); ANT I1 with 2 member of Yosii’s (1969) “formosana group”. It can be
dorsal and 1 ventral cup sensilla; ANT 111 not subseg- separated from formosana by having circumanal setae H
mented, with cup sensilla and distal sensilla exposed, spine-like and M’ blunt, spine-like; solomonensis and
lying in depressions; ANT IV not subsegmented. Outer modesta have the same circumanal setal pattern as longi-
maxillary lobe consists of simple palp and 1 sublobal hair digita, separation is possible on the basis of color pattern
(Fig. 241). Labrum not seen. Dorsal setae of head short, and shape of the claws. The Hawaiian species that come
fine; 5 unpaired facial setae (Fig. 242). closest to longidigita are hrevifihra, sylvestrutilis and tesse-
Foreleg. Coxa, trochanter and femur not clear or ob- lata. Separation of the 4 species can be accomplished
scured in preparation; tibiotarsus with 3 cup sensilla and 1 using the charwteristics listed in Table V.
(visible) “courte epine” on anterior surface (Fig. 243),
posterior surface without “courte epine” (Fig. 244); pre-
tarsus with anterior and posterior setulae; unguiculus Dicyrtoma (Calvatomina)bellingeri sp.n. (Figs 257-289)
with pseudonychia, tunica, up to 2 outer teeth and 2 inner
teeth; unguiculus with 2 corner teeth, subapical filament Maferial examined. Holotype ( 9 ) and 3 paratypes prcscrvcd in
ethanol, 2 paratypes o n slides. Hawaii, Kauai County, Kauai, Na Pali
heavy, tapering, longer than unguiculus (Fig. 245). Wilderness Area, Hanakoa Campsite, forest, near stream, sifting litter,
Mesoleg. Coxa, trochanter, and femur not clear or 22 March 1986, K . A. Christianscn.
obscured in preparation; tibiotarsus with 5 cup sensilla
and 2 “courtes epines” on anterior surface (Fig. 246), Description
posterior surface without visible “courte epine” (Fig.
247); pretarsus with anterior and posterior setulae; unguis Color. Background opaque to white with brown-purple
with pseudonychia, tunica, outer tooth and 2 inner teeth; pigment laid down in irregular polygons. Antenna distally
unguiculus with strong corner tooth and small second dark purple-blue, becoming lighter basally. Vertex of
tooth, subapical filament heavy, tapering, longer than head with light purple macula between eyepatches and
unguiculus (Fig. 248). flanked by smaller, darker maculae. Frons with purple
Metaleg. Coxa, trochanter and femur not clear or mosaics forming an irregular band extended to postgenal
obscured; tibiotarsus with 5 cup sensilla and 3 “courtes area. Lower frons with medial macula, frontal oral area
epines” on anterior surface (Fig. 249), no visible “courte bordered with purple mosaics. Body with purple lateral
Zoologicu Scnptu 19
 Dicyrtomidae (Collembola) of Hawuii 91

Figs lY4-22.5. Dicyrroma (Calvutornina)tesselatu sp.n.-lY4. Left eyepatch.-lY5. Antenna1 segment I.-lY6. Outcr maxillary lobc.-l97. Facial
sctae.--198. Forccoxa.--IYY. Foretrochanter.-200. Forefemur.-201. Foretibiotarsus, anterior surface.-202. Forctibiotarsus, posterior
surfacc.-203. Foreclaw.-204. Mesocoxa.-205. Mcsotrochantcr.-2W. Mesofemur.-207. Mcsotibiotarsus, anterior surface.-208. Mesotibio-
tarsus. posterior surface.-209. Mesoclaw.-210. Metacoxa.-2f l . Metatrochanter.-212. Metafcmur.-2I3. Metatibiotarsus, anterior surface.-
214. Metatibiotarsus. posterior-surface.-2l5. Metaclaw.-2f6. Collophore.-2f 7.Corpus of retinaculum.-218. Manubrium. lateral view.-219.
Dens, ventral surface.-220. Dens, dorsal surface.-22/. Dental setae E,-E,.-222. Mucro.-223. Anal papilla, female.-224. Female subanal
appcndage.-225. Parafurcular sctae.
Zoologica Scripta IY
02 R. J . Snider

Frg. 2.?X llicyrtonia (C‘ulvrrrorninu) loorigirligirci sp.n.. habitus, lateral

vicw. Total length up to 2 mm

(Fig. 265), posterior surface with 1 “courtc epine” (Fig.

band becoming darker posteriorly; a large purple V 266); pretarsus with anterior and posterior setulae; unguis
postero-dorsally and pair o f light purple rectangular with pseudonychia, tunica, outer tooth, and 2 inner teeth;
patches. Upper and lower valves of anal papilla with unguiculus with sharp, straight corner tooth, subapical
ventral purple spots. Legs from lower femur distally with filament over 1/2 length of inner edge of unguis, tapering
purple-blue, collophore and furcula with light purple beyond tip of unguis (Fig. 267).
dusting (Figs 257, 258). Mcsoleg. Coxa with 3 anterior setae and 1 “courte
epine” (Fig. 268); trochanter with 3 anterior and 1 pos-
+
Morphology. Eyes 8 8, ocelli subequal. Mean antenna1 terior setae (Fig. 269; femur with 9 anterior and 3 pos-
ratio 4 : 1.5 : 18 : 5 ; ANT I with 5 dorsal and 2 ventral sctae; terior setae, outer cup sensillum, 2 posterior setulae, 1
ANT I1 with 2 dorsal and 1 ventral cup sensilla; ANT Ill anterior “courte epine” and basal posterior oval organ
not subsegmented, armed with cup sensilla, subapical (Fig. 270); tibiotarsus with 5 cup sensilla and 3 “courtes
sensilla exposed, lying in shallow depressions; ANT IV epines” o n anterior surface (Fig. 271), 1 “courte Cpine”
not subsegmented. Outer maxillary lobe consists of on posterior surface (Fig. 272); pretarsus with anterior
simple palp and 1 sublobal hair (Fig. 2.59). Labrum with and posterior setulae; unguis with pseudonychia, tunica,
setal pattern 6/S, 5,4. Dorsal setae of head reduced, small outer tooth and 2 inner teeth; unguiculus with 2 corner
(Fig. 260); 6 unpaired facial setae (Fig. 261). teeth and small subapical tooth, subapical filament short,
Foreleg. Coxa with 1 anterior seta (Fig. 262); tro- about even with tip of unguis (Fig. 273).
chanter with 3 anterior and 1 posterior sctae (Fig. 263); Metaleg. Coxa with 4 anterior setae and 1 “courte
femur with 5 anterior and 5 posterior setae, 2 posterior Cpine” (Fig. 274): trochanter with 4 anterior and 1 pos-
setulae, outer cup sensillurn, anterior “courte epine” and terior setae (Fig. 275); femur with 10 anterior and 1
basal posterior oval organ (Fig. 264); tibiotarsus with 4 posterior setae, outer cup sensillum. 2 setulae, 1 anterior
cup sensilla and 3 “courtes epines” on anterior surface “courte epine” and 1 posterior basal oval organ (Fig.

231

235
232 233 234 237
Figs 227-237. Ilicyrtorna (Cblvutorninu) rnirrodcnratu sp.n.-227. Left eycpatch.-228. Foreclaw.-22Y. Mctaclaw.-230. Collophore. lateral
vicw.-2SI. Corpus of rctinaculum.-232. Manubrium, lateral view.-2.?3. Dental setac E,-E3.-234. Mucro.-235. Anal papilla, female.-236.
Female subanal appendage.-237. Parafurcular setae.

Zoologiro Srripru 14,

 Dicyrtomiclae (Collemholu) of Huwaii 93

\
239 240

,\

- 6
c

’
---==
/#+-

Figs 239-2S6. Dicyrioma (Culvaiomina) longidigiia sp.n.-239. Right eyepatch.-240. Antcnnal segment 1.-24/. Outer maxillary lobc.-242.
Facial sctac.-243. Foretibiotarsus. anterior surfacc.-244. Forctibiotarsus, posterior surface.-245. Forcclaw.-246. Mesotibiotarsus, anterior
surfacc.-247. Mcsotibiotarsus. posterior surface.-248. Mesoclaw.-249. Mctatibiotarsus, anterior surface.-250. Mctatibiotarsus, posterior
aurfacc.-251. Metaclaw.-252. Mucro.-253. Anal papilla. femalc.-254. Fcmalc subanal appendagc.-255. Parafurcular sctae.-2.56. Dorsal
setae of abdomen.

276); tibiotarsus with 5 cup sensilla, 3 “courtes @pines”on and 2 inner teeth; unguiculus with 2 corner teeth, subapi-
anterior surface (Fig. 277), 1 “courte dpine” o n posterior cal filament short, fine, reaching tip of unguis, outer
surface (Fig. 278); pretarsus with anterior and posterior lamella expanded greater than meso-unguiculus (Fig.
setulae; unguis with pseudonychia, tunica, outer tooth, 279).
+
Great abdomen. Collophore with 1 1 subapical and
+
1 1 lateral setae, sacs warty (Fig. 280). Corpus of
tenaculum with 4 setae, ramus with 3 teeth and basal horn
(Fig. 281). Manubrium with 11 + 11 dorsal setae (Fig.
282). Dens with 4 , 2 , 1 , 1 . . . 1 Ve setae (Fig. 283), dorsal
E and L setae smooth DI-D3 subequal (Fig. 284), EI/E2
= 1.20, E3/E2 = 1.25 (Fig. 285). Mucro with numerous,
regularly spaced inner and outer teeth (Fig. 286). Circu-
manal setae follow the pattern
M N A(, T H G and L M’ N’
+ + - - + f . + + +
F i p 2S7. 258. Dicyriornu (Calvaiorninn) bellingeri sp.n.-257. Habitus,
lateral view.-25#. Habitus. dorsal view. Total lcngth up to 0.85 mm. (Fig. 287); female subanal appendage heavy, sharp,
Zoulrigicn Scripla 19
94 R . J . Snider

I 1 263
:’ B Q
261
260

264

267 ‘I

_--- 273
-
271 272
279

280
282
\!
28 1 -’,--- Y’
I

83
.... . . - - -
277 278
r

/” 285 286 288

Figs 25Y-289. Dicyrtoma (Calvatomina) bellingeri sp.n.-259. Outcr maxillary lobe.-260. Dorsal setae of head.-261. Unpaired facial sctae.-
262. Forccoxa.-263. Foretrochanter.-264. Forefemur.-265. Foretibiotarsus, anterior surface.-266. Forctibiotarsus, posterior surface.-267.
Foreclaw.-268. Mesocoxa.-269. Mcsotrochanter.-270. Mesofemur.-271. Mesotibiotarsus, antcrior surface.-272. Mesotibiotarsus, posterior
surface.-273. Mesoclaw.-274. Metacoxa.-275. Metatrochanter.-276. Mctafcmur.-277. Metatibiotarsus, anterior surface.-278. Mctatibio-
tarsus, posterior surface.-279. Metaclaw.-280. Collophorc.-281. Corpus of retinaculum.-282. Manubrium, lateral vicw.-283. Dens, ventral
surface.-284. Dens, dorsal surface.-285. Dental setae EI-E3.-286. Mucro.-287. Anal papilla, fcmalc.-288. Parafurcular setae.-289.
Posterior abdominal setae.

Zoologica Scripta I9
 Dicyrtomidae (Collembola) of Hawaii 95

tapered and curved at apex. Parafurcular setal pattern 3 & Weiner (1987) have presented differences between
spine-like and 2 setaceous (Fig. 288). Bothriothrix A, B, number, shape and pattern of dorsal setae found on the
C normal, D missing. Body setae fine in thoracic region, great abdomen which clearly separate Ptenothrix and
becoming spine-like posteriorly (Fig. 289). Length up to Bothriovulsus.
0.85 mm. During this study, differences between Ptenothrix spp.
and Papirioides spp. appeared so slight that at the sugges-
Diagnosis. Dicyrtoma (Calvatomina) hellingeri sp.n. is a tion of Betsch (in lift.), Papirioides was reduced to a
member of Yosii”s (1969) “formosana group”. It can subgenus of Ptenothrix. Ptenothrix (Ptenothrix)hawaiien-
easily be separated from those published species on the sis sp.n. exhibited the characteristic pattern of 1 , 1 blunt,
basis of circumanal setae pattern and claw morphology. unpaired, median facial setae. This has been suggested as
Among members of the Hawaiian fauna, bellingeri has a indicative of Holotropical Ptenothrix spp. (Yosii 1969;
unique color pattern that will distinguish it from all Betsch 1980) and used by Christiansen & Bellinger (1981)
others. The circumanal setal pattern is similar to syfvestra- for North American identification of the genus.
tilis, however, the parafurcular setal pattern is S/O for that However, contrary to Christiansen and Bellinger,
species and 312 for hellingeri. Snider (19850, h ) has shown that two species described
from the southeastern United States had facial setal
patterns composed of seta-like elements in the arrange-
Comments on systematics ment of: 1, 1, 1 , 2, 1 , 1, 1, 3 for Ptenothrix (Ptenothrix)
renateae (Snider) and 1 , 1, 1, 1, 1, 1, 3, 4 for Ptenothrix
The tribe Ptenothricini defined by Richards (1968) is (Ptenothrix) castanea (Snider). Likewise, Ptenothrix (Pte-
cornposed of genera exhibiting bothriothrix D and two nothrix)vittuta (Folsom) from North America and Japan
differentiated setac on tibiotarsus P?. The Hawaiian has 1, 1, 1 , 2 , 1, 1, 1, 3 setae similar to renuteae.
fauna consists of two subgenera represented by: Ptenoth- Among the Hawaiian species Ptenothrix (Papirioides)
rix (Ptenothrix) hawaiiensis sp. n., Pteraothrix (Papir- kauaiensis sp.n. had facial setae in the pattern: 1, 1 , 2 , 2 ,
ioides) kauaierisis sp.n., serrata sp.n. and duhia. Besides 1 , 1 , 2, 3 with the first two setae blunt, similar to P.
the characteristics cited above, Richards (1968) used the hawaiiensis; while Ptenothrix (Papirioides) serrata had
presence or absence of subannulation of antenna1 seg- the pattern: 1, 1, 1 , 2, 1, 1, I , 2 with all setae similar.
ments 111 and IV in his diagnosis. Betsch (1980) dis- Unlcss more new convincing morphological character-
counted this feature. H e found that subannulation istics are found, the presence or absence of blunt,
appeared in stored specimens of Ptenothrix, Dicyrtoma unpaired facial setae becomes doubtful for absolute gen-
and Dicyrtomina. eric separation.
Richards ( 1968) separated Ptenothrix and Papirioides The relationship of Papirioides to Ptenothrix is very
from Bothriovulsus sp. by the presence of bothriothrix A. close. They share almost all morphological features. This
This appears to be a significant characteristic for division problem was recognized by Stach (1465) and Richards
of Ptenothrix from Bothriovulsus. More recently Betsch (1968). However, the abdominal dorsal posterior pro-

290

Figs 290-2’93. Comparison of Prrnothrix (Papirioides) spp. with abdominal protuberances and Pfenorhrix (Ptenothrix) hawaiiensis sp.n.-2W.
Ptcwofkrix (Papirioides)duhiu. abdominal protuberance .-2YI. Ptenothrix (Pupirioides)serrufu, abdominal protuberance, dorsal view, collapsed.-
2Y2. Prcviotlirix (Papirioides) kuuairnsis, abdominal protuberance.-2YS. Pfenothrix (Pupirioides) hawuiwnsis, dorsal view of posterior abdomen.
Arrows indicate setae associated with abdominal protUbCrdnce.
96 R . J . Snider

and serruta (Fig. 296) are extremely variable and easily

intergrade with Ptenothrix hnwaiiensis (Fig. 297).
It seems logical, until more species are described, that
Pupirioides should be positioned as a subgenus of Pte-
nothrix. In addition, here Dicyrtomu has been applied to
members of the tribe Dicyrtomini as the generic name.

4
Thus Dicyrtornina, Gihheruthrix and Calvatominu be-
come subgeneric designations. To separate them as gen-

= era based on tunica inflated or not, and development of

f
the pseudonychia, seems questionable at best.

Zoogeographic considerations

World-wide Ptrnothrix sensu stricto is represented by 58

294 295 296 297 published species. Of these, 19 may be classified as
I.i‘gs 2Y3-297. Comparison o f female upper and Iowcr anal valvc glands Holotropical (Yosii 1969; Betsch 1980). Yosii (1969)
found among H iiwai i a n Pten oihrix ( Pupirioitlrs) spp. and I’rcriothrix suggested that we could recognize Ptenotlzrix from the
(f‘tetiotlirix) huwuiieri.si.s sp.11.--2Y4. Ptenorhrix (Pupirioides) tiuhicr, tropics by the presence of two blunt, unpaired facial sctae
upper and lower valve glands of female .-XiPtenotiirix
. (f‘upirioitles)
krnraiensis. upper and lower glands.-Z96. f‘tef7otkri.u (PcqJirioides) and a blunt anal seta A,,. Ptenothrix (Ptenothrix) haw-
scrru/u. upper and lowcr glands.-297. Prenorhrix (f‘teriothrix) huw- aiiensis fits into this category. However, we should be
uiiensis, uppcr and lower glands. careful in generalizing from these characteristics. Betsch
(1980) saw the possibility of splitting the Holarctic and
Holotropical species into distinct genera. Table VI
tuberance seems to make separation of the two subgenera appears to support, at least partially, this concept with
possible. Even though both P . (Pupirioides) kauaiensis reservations about Ptenothrix (Ptenothrix)atra, marmor-
and serrata share absence of anal seta G with the “ciliata” utu, and vittuta.
group of Ptenothrix (Yosii & Lee, 1963), it continues to The distribution of P. utrn is Holarctic with a southern
make sense to use subgeneric designations. range reaching Costa Rica (Denis 1931, 1933). This
Within the Dicyrtomini there exists a similar situation species has both seta-like unpaired facial and A(, anal
where a “group” or subgenus, depending on interpret- setae. L,ikewise, P . mnrmorutu is Nearctic and exhibits
ation, exhibits an abdominal protuberance similar to the two unpaired, blunt facial setae, while A(, is seta-like. Its
condition found in Ptenothrix (Papirioides) spp. Of the range is south to Mexico (D.F.) which places it in the
three reported species in the “Pseudodicyrtominu group”, Neotropical region (Handschin 1928). In addition, mur-
Dicyrtoma (Calvatominu) homhuyensis Yosii ( 1966), morotu was recorded from Japan (Uchida 1Y52h, 1954,
Dicyrtoma (Calvntomina) truknnn (Uchida), and Dicyr- 1955, 1959). Another species, P. vittata, is found in North
toma (Calvntomina) verrucosu (Handschin), all appear to America (Folsom 1896; Christiansen & Bellinger 1981),
have spine-like setae associated with the protuberance in Japan (Yosii & Lee 1963), North Vietnam (Stach 1965)
a like manner to P . (Papirioides) duhia. and Australia (fasciafa Womersley 1932, 1939; Yosii &
Comparison of kaziaiensis and serrata with P. (Pnpir- Lee 1963). Ptenothrix vittutu has seta-like unpaired facial
i d e s ) duhiu (Fig. 290) shows that both (Figs 291, 292) setae and a blunt A. seta.
exhibit a pattern of spine-like setae on the posterior of the It appears that both marmorata and virtara exhibit
great abdomen in the location of the protuberance. This characteristics of “groups” found in the Holarctic and
feature is lacking in P. (Ptenorhrix) hawuiiensis (Fig. 293). Holotropical realms. Further collections must be made or
Recently, Betsch ( i n l i l t . ) has suggested that the anal examined from the tropics before separation of groups
glands of female Ptenothrix spp. and Papirioides spp. can be made with absolute confidence.
might be useful for subgeneric or even generic separation In the diagnosis presented here, P . hawaiiensis is
(Delamare & Massoud 1963; Arle 1976; Soto-Adames placed close to ciliophora and saxatilis based on mor-
1988). However, comparison of these glands showed that phology of facial, labral and anal valve setae. However,
P. (Papirioides) duhiu (Fig. 294), kuuaiensis (Fig. 295) both the latter species are endemic to Korea and do not

Tablc VI. Ptcnothrix (Ptcnothrix) spp. listed by region of the Holorropic realm. Note rhuf A& = uriul .sera
A,) is blunt. AoS = setuceous und R = 2 bhmt, utipcured fuciul setue. S =facial serae sharp, setu-like

tlolotropic
Neotropical Oriental Australian Ethiopian
~~ ~ ~ ... ~~~~~~~ ~~ ~~ ~.
..
~ ~ -. . . . . ..

argentina A,,b,B fiscelluiu? grucillicornis Al,b,R tnurphyi A&,B

utru A,,s,S gib ~ O S UA,,b,B hawuiiensis A,,b,B i~ioluceopusA,,[>,B
horinrunu A,,b.B himaluyensis A,,b,B 1)ittatu AI1b.S violucc~a?
h rusilirvisis A,,b ,B kerulrre A,,b,B
tloniinicunu A&, B pula wunensis A&, R
niurrnorutu A,,s.B puldiellrc,s?
mritrgur’ A,,b,B

Zoologica Scripu I 9
 Dicyrtomiriue (Collemholu) of Hawuii 97

Table V1 I . llistrihrttion by region of’ Ptenothrix (Papirioidcs) spp. l’ablc VIII. Ptcnothrix (Papirioidcs) . s ~ / J/runtul
. setulpurferns of’heud

Regions within realms Pattern

Eastcrn Palcarctic Oriental A us t ral i a n
-- ~~ __ _. ~ ~. -- ~

1-1-2-2-1-1-1
urc~rtirr~hc~rc.rrl/rr jacohsoni duhiu I-I-1-1-2-1-1
j a w hs on i kuuuiensis 1-1 -2-1- 1-1 -1 ( 3 )
nrirnbilk serrutu 1-1-1-1-1-1-1
rriwii Unknown
I-1-1-2-1-1-1
Unknown

have only 2 blunt, unpaired facial setae. Jt is also sugges-

ted that t‘.gracillicornis is close to huwuiiensis. If vittutu is
excluded as only recently being transported to the Austra- nounced abdominal protuberance. It appears that duhia
lian region, then the closest regionally endemic allied and serrufa are more closely allied to each other than they
species to Iiii wuiiensis becomes grucillicornis. are to kauaieizsis. Betsch (1980) reports a species (yet
Thc species jucohsoni and uerioi are both Palearctic in undescribed) close to duhia from Madagascar and
origin (Table VII). The abdominal protuberance is pro- suggests it may represent an import.
nounced, unlike the low structure seen o n aequituhercu- Members of Dicyrtoma (Calvatomina) s p p . are pre-
lutn and duhia. The heavy, unpaired facial setae are dominant in the Holotropical realm with 12 of 25 species
similar in position to those found in Pteizothrix spp. found in the Australian region. It is apparent that Culva-
However, they are not blunt and o f uniform diameter as fominu has successfully established in the Oriental-
those seen in species like P . ~ t i n g a e grucillicornis,
, or Australian regions. Today only six species are found in
murphyi. The unpaired facial setae of Pupirioidrs spp. the Holarctic realm (Table IX).
taper in at least the last 1/8 of their length to a point, The species D. (Calvutomina) madestris sp.n. and mic-
giving a gladiform shape. All species, so far, exhibit this rodentutu sp.n. clearly fall within the “rufesceizs group”.
shape. The Pacific species unpaired facial setal patterns They both exhibit spine-like parafurcular setae and have
are presented in ’Table VIII. M’ reduced to a small seta. They are the first species of the
From the known distributions, the origins of Pupir- “rufesceizs group” to be recorded from the Australian
ioidrs would seem to be eastern Palearctic with dispersion region and were taken only on Kauai.
through the Oriental region to the Australian region. Of The “formosnna group” species are very similar in
the three Hawaiian species, duhiu has the most pro- many features, especially with 3/2 parafurcular setae. An

Table IX. Known di.strihrrtions of Dicyrtoma (Calvatomina) spp.

India
Japan
Philippines

Solomon la.
Solomon Is.
Surinam
Solomon Is.

India
Hawaiian Is.
Hawaiian Is.
Angold
(U.S.A.)
Gambia
India, Burma. Sri Lanka
India
Sudan
Japan (Europe) Pucrto Rico
India
0kin a w a Java

Rhodcs
Hawaiian I s .
Hawaiian Is.
‘Taiwan
Hawaiian Is.
Solomon Is.
Pucrto Rico
Solomon Is.
Hawaiian Is.
Hawaiian Is.

Zoologicu Scripa 19
98 R. J . Snider

exception is sylvestratilis with S / O . This species resembles ington produced the hahitus drawings for Dicyrronur (Cu/vatoniinu)
solomonensis and rnodestu from the Soloman Islands. Rrevrfihru n.sp. Many thanks arc offered to Dr J.-M. Bctsch for hclpful
suggestions about systematics and zoogcography.
The Hawaiian D. (Calvatominu) spp. appear to be
endemic to specific islands with a possible exception of
hrevifihru which was found on the islands of Hawaii,
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Zvolofiica Scriptu I 9
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Zoologicu Scriptu I9