Research Article |
Corresponding author: Cristian C. Lucanas ( cclucanas@up.edu.ph ) Academic editor: Oana Teodora Moldovan
© 2016 Cristian C. Lucanas, Ireneo L. Lit Jr..
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lucañas CC, Lit IL Jr. (2016) Cockroaches (Insecta, Blattodea) from caves of Polillo Island (Philippines), with description of a new species. Subterranean Biology 19: 51-64. https://doi.org/10.3897/subtbiol.19.9804
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Cockroaches collected during a protracted series of fieldwork in several limestone caves in Polillo Island, Philippines were taxonomically studied. A new species of Nocticola (Nocticolidae), N. gonzalezi Lucañas & Lit, sp. n., is described. The male of Periplaneta banksi Hanitsch (Blattidae) is described for the first time. Altogether, the present cave cockroach fauna of the island consists of six species.
Blaberidae , Blattidae , Burdeos, cave cockroaches, Ectobiidae , Nocticolidae , troglobiont
Cockroaches are well represented in cave ecosystems. They are classified as: trogloxene, troglophile, guanobyte or troglobiont depending on their specialization or adaptation to the cave environment (
Unfortunately, very few studies have been conducted regarding the diversity of Philippine cockroaches, maybe because the group includes species that are considered household pests. Thus, limited information is available on the country’s fauna, especially in cave ecosystems. In recent years, the increasing interest in cave studies in the country has provided an opportunity to study the diversity of cockroaches in Philippine limestone caves.
The first documented troglobiont cockroaches in the Philippines were species of Nocticolidae from Rizal Province (Bolivar 1982), a few kilometers to the east of what is now Metro Manila. These were: Nocticola simoni Bolívar, 1892 (from a cave in San Mateo town), and Nocticola caeca Bolívar, 1892 (from Cueva de Talbac, in what is now Antipolo City).
Unfortunately, due to the limited knowledge on Philippine cockroach diversity, many species have often been misidentified.
Other than those, there have been very few studies regarding the diversity of cockroaches in caves. This paper provides a list of cockroaches collected from several caves in Polillo Island, Quezon Province in the Philippines.
All examined specimens were either pinned (for larger specimens) or preserved in vials of 95% ethyl alcohol, unless otherwise indicated as mounted on slide using Canada Balsam. Slide preparations were made starting with cold maceration by soaking in 10% aqueous Potassium Hydroxide (KOH) solution for 24 hours at ambient room temperature. They were then washed in distilled water to remove unwanted undissolved internal contents, and then run through graded alcohol series and then anhydrous clove oil to remove excess ethanol. Finally, they were mounted on microscope slides using Canada Balsam. The following parameters were measured using a calibration micrometer for pinned specimens and calibrated ocular micrometer for slide mounted specimens:
TL (Total length) the length from the tip of the vertex (if exposed) or pronotum up to the tip of the tegmina at rest;
BL (Body length) the length from the tip of the vertex (if exposed) or pronotum up to the tip of the subgenital plate (in case of brachypterous/micropterous/apterous species, total length = body length);
Pn Pronotum length × width along the longest axis;
Tg Tegmina length along the longest axis.
Standard deviations for each measurement were also noted. Terminologies used for male genitalia follow Klass (1977), and
Illustrations were made by tracing printed photomicrographs of clear slide-mounted specimens. The degree of sclerotization and pigmentation was made through line thickness and by stippling. All specimens are deposited in the Entomological Collection of the UPLB-MNH.
Holotype: male, slide-mounted. Original label: “Philippines: Polillo Island (Quezon Province): Cave 5, Sitio Puting Bato, Barangay Aluyon, Burdeos, 13.v.2013 CC Lucañas” “UPLBMNH BLA-00220”, “HOLOTYPE / Nocticola gonzalezi Lucañas & Lit [laser printed on red slide label]”
Paratypes: 1 male, 1 female, 5 nymphs, same locality as holotype but 10.x.2013: (CC Lucañas, UPLBMNH BLA-00221, ♀ (slide); -00222, ♂; -00223, nymph (slide); -00224a-d, 4 nymphs). “PARATYPE / Nocticola gonzalezi Lucañas & Lit [laser printed on blue slide label or handwritten on blue label]”.
Philippines: Polillo Island (Quezon Province): Cave 5, Sitio Puting Bato, Barangay Aluyon, Burdeos, on calcite rubble with insectivorous bat guano.
Male brachypterous; female apterous. Compound eyes reduced, much smaller in female. Femur Type C1. Tarsi simple, symmetrical. Tergal gland absent (simoni-species group). Accessory hook-like phallomere (L2d) dagger-like.
This species can be distinguished from other troglobiotic Nocticola by the presence of an accessory hook-like phallomere which was only reported from two epigean species, N. scytala Andersen & Kjærandsen and N. clavata Andersen & Kjærandsen from Ghana, West Africa (
This species is similar to N. simoni and N. caeca in terms of coloration but differs in terms of size, N. gonzalezi being smaller. Unfortunately, the male genitalia of N. simoni and the male of N. caeca was not described by
Many Nocticola species are narrow endemics even among caves within a contiguous area (e.g. N. simoni was collected from a cave in San Mateo, Rizal Province and N. caeca from nearby Antipolo, also in Rizal Province). Considering this, as well as the morphological characteristics and the isolation of Cave 5 in Puting Bato, Burdeos, Polillo Island, we iterate the distinctness of N. gonzalezi.
Size (mm): ♂, TL: 2.36 ± 0.13 mm; Pn: 0.50 ± 0.10 × 0.93 ± 0.05 mm; Tg: 1.13 ± 0.2. ♀ TL: 2.5 mm; Pn: 0.50 × 0.95 mm.
Male. (Fig.
Nocticola gonzalezi Lucañas & Lit, sp. n.: A Adult male B adult female C male head, lateral aspect D Prothoracic leg E male genitalia: (L1-3: sclerites of left phallomere; L2d: L2 dorsal; L2v: L2 ventral; R1-3: sclerites of right phallomere; vph: ventral phallomere) F female subgenital plate G male and H female habitus.
Female. (Fig.
Nymphs. Similar to female. Opaque white. Eyes reduced to 3-4 distinct facets.
The species is named after Dr. Juan Carlos T. Gonzalez, the current director of the UPLB-MNH, in recognition of his pioneering efforts toward biodiversity documentation and conservation in the Polillos, a group of islands, east of Luzon and politically a part of Quezon Province.
Philippines: Polillo Island.
Collected only from a single cave in Burdeos, Polillo Island, Quezon. The cave is deposited with porous calcite rocks and thin guano deposits from periodical roost of insectivorous bats, Hipposideros coronatus (Peters, 1871) and Miniopterus schreibersii (Kuhl, 1817). The site is relatively dry and rocky compared to the damp and sticky clay that is found throughout the rest of the cave, especially during the rainy season. They are quite fast and usually hide among the holes in the calcite rocks. They are found among and probably serve as prey for nymphs of whipspiders (Arachnida: Amblypygi), whipscorpions (Thelyphonida) and adult short-tailed whipscorpion (Schizomida).
Due to the narrow distribution of this species, it is suggested that the species be considered “Vulnerable” and that the cave from which it was collected be protected from further anthropogenic disturbances.
Because of its morphological characteristics, this species can be classified as a true cave-dweller or troglobiotic.
Periplaneta
banksi
:
8 males, 10 females, 3 nymphs: Philippines: Polillo Island (Quezon Province): Bulalon Cave, Poblacion, Burdeos, 23.ii.2013 (IL Lit, Jr/ OL Eusebio/ AR Larona, UPLBMNH BLA-00232-00234, ♂, -00235 ♀); 25.v.2007 (IL Lit, Jr/ OL Eusebio/ AR Larona, UPLBMNH BLA-00236-00242, ♂, -00243-00245, ♀, -00246-00248, nymphs); 11.vi.2008 (OL Eusebio, UPLBMNH BLA-00528-00531, ♀).
Reddish-brown throughout. Fore femur A2 (Fig.
Periplaneta banksi Hanitsch, 1931: A Adult female B head C prothoracic leg D tarsal claws E male genitalia: (L1-3: left phallomere sclerites; L2v: L2 ventral sclerite; R1-3: right phallomere sclerite; vph: ventral phallomere) F L2v close-up G male supra-anal plate H female subgenital plate.
Size (mm): ♂ TL: 45.71 ± 2.29; BL: 37.42 ± 3.69; Pn: 11.57 ± 0.79 × 14.86 ± 1.46; Tg: 35.71 ± 2.29; ♀ TL: 48.5 ± 2.01; BL: 41 ± 2.53; Pn: 11.6 ± 0.51 × 15.8 ± 1.03; Tg: 37.7 ± 2.41.
Male. Reddish-brown throughout. Vertex exposed. Ocelli present. Pronotum wider than long, markings indistinct. Tegmina well-developed, extending beyond abdomen. Fore femur A2. Pulvilli present on four proximal tarsi. Minute sub-apical spine present on three proximal tarsi. Mid- and hind tibia with several minute spines on outer aspect. Hind basitarsus longer than succeeding tarsi combined. Arolia minute. Tarsal claws simple, symmetrical. Abdominal tergal gland absent. Supraanal plate testaceous, deeply divided, each lobe rounded with minute round projection. Genitalia as illustrated, genital hook angulate, with minute sub-apical process.
Female. Similar to male except that supra-anal plate rounded with deep division. Subgenital plate valvular.
Philippines, Taiwan
This is the first record of the species inside caves. The species is abundant and most individuals hid among the crevices of the cave walls. They were found from the twilight to the dark zone. Nymphs and mating pairs were often seen, suggesting that the species can complete their life cycle within the cave. Despite that, P. banksi is classified as troglophilic because of the lack of distinct troglobiotic characters and the possibility of the species surviving outside cave conditions.
Both nymphs and adults are possibly preyed upon by whipspiders, Charon sp., and huntsman spiders, Heteropoda sp.
The species resembles P. lata (Herbst, 1786), but differs primarily by its larger size.
Leucophaea
striata
Kirby, 1903: 378 (Lectotype: ♂, Batu Cave, Selangor, Malaysia, BMNH, designated by Princis 1956);
Pycnoscelus
striatus
,
22 males, 15 females, 38 nymphs: Philippines: Polillo Island (Quezon Province): Bulalon Cave, Poblacion, Burdeos, v.2010 (JMA Encinares, UPLBMNH BLA-00443-00455 and -00500-00505, ♂; -00456-00466, ♀; -00467-00497, nymphs), 24-27.ii.2007 (IL Lit, Jr/ OL Eusebio, UPLBMNH BLA-00507-00509, ♀); Cave 2, Sitio Puting Bato, Barangay Aluyon, Burdeos, 23.ii.2013 (CC Lucañas, UPLBMNH BLA-00517, ♂; -00518, ♀), 23.v.2013 (CC Lucañas, UPLBMNH BLA-00524 ♂; -00522-00523, nymphs); Cave 3-4, Sitio Puting Bato, Barangay Aluyon, Burdeos, 23.ii.2013 (CC Lucañas, UPLBMNH BLA-00511, -00515, ♂; -00512, -00516, ♀; -00513-00514, nymphs); Cave 5, Sitio Puting Bato, Barangay Aluyon, Burdeos, 24.ii.2013 (CC Lucañas, UPLBMNH BLA-00519-00521, nymphs).
Size (mm): ♂ TL: 14.25 ± 1.32; BL: 13.59 ± 1.15; Pn: 4.57 ± 0.42 × 5.91 ± 0.65; Tg: 9.68 ± 1.56; ♀ TL: 21.39 ± 1.91; BL: 17.67 ± 1.97; Pn: 5.97 ± 0.40 × 7.42 ± 0.43; Tg: 16.67 ± 1.91.
Male. Often less than half the size of females. Vertex exposed. Ocelli present. Pronotum rhomboid, almost as long as wide, reddish-brown, densely punctured. Tegmina variable, reaching slightly beyond abdomen or reaching only up to 7th abdominal tergite. Fore femur C1. Mid- and hind femur unarmed. Pulvilli present on four proximal tarsomeres. Hind metatarsomere slightly longer than succeeding tarsomeres combined. Tarsal claws simple, symmetrical. Arolia minute. Abdominal tergites unspecialized. Supraanal plate entire, testaceous. Genitalia as illustrated. Left style absent. Right style ovoid, rounded apically. Genital hook on right side.
Female. Almost similar to males except larger and robust. Wing size highly variable. Subgenital plate entire.
Malaysia (Malacca), Indonesia (Sumatra, Kalimantan), Philippines.
The foregoing description of the right style (i.e. rounded apically) differs from that of
3 females: Philippines; Polillo Island (Quezon Province): Cave 2, Sitio Puting Bato, Barangay Aluyon, Burdeos, 23.v.2013 (CC Lucañas, UPLBMNH BLA-00525-00526), v.2010 (JMA Encinares, UPLBMNH BLA-00527).
Size (mm): ♀ TL: 21.5 ± 0.71; BL: 18.5 ± 0.71; Pn: 6.5 ± 0.71 × 8.5 ± 0.71; Tg: 17.5 ± 0.71.
Male: Unknown
Female: Body generally convex. Vertex concealed. Ocelli present. Pronotum medially wider than long, black throughout except for yellow anterolateral markings, minutely punctate. Tegmina well-developed extending beyond abdomen, hyaline black-brown. Fore femur C1. Hind basitarsus longer than succeeding tarsal segments combined. Pulvilli present on four proximal tarsomeres. Arolium large. Tarsal claws symmetrical. Abdomen medially brown, darkening distally.
The species was collected on thick guano piles of mix bat populations in Cave 2. In contrast with P. striatus, the species was present only on the top layer of the guano and does not burrow much even if disturbed.
This species or form is closely similar to P. striatus and may prove to be only a variation. For now, it is treated separately because of its darker coloration, robust abdomen, rather convex body, large arolia and the presence of narrow anterolateral band on its pronotum. The absence of a male specimen hinders further identification.
1 female: Philippines: Polillo Island (Quezon Province): Cave 3-4, Sitio Puting Bato, Barangay Aluyon, Burdeos, 01.iii.2013 (CC Lucañas, UPLBMNH BLA-00533).
Size (mm): TL: 13.50; Pn: 3.00 × 4.00; Tg: 10.00.
Male: Unknown.
Female: Vertex of head slightly exposed. Ocelli minute. Pronotum anteriorly ellipsoid, wider than long, with brown central maculae, anterior and lateral areas hyaline. Fore femur type A2. Mid femora armed with row of spines on outer aspect. Hind femora missing from specimen. All tarsomeres with pulvilli, 1-3 spined. Tarsal claws symmetrical with minute dentations. Arolium present. Tegmina well-developed.
Accidentals. Since only one female specimen was collected and troglomorphic characters are absent, it can be said that its presence in the cave was accidental.
3 nymphs: Philippines: Polillo Island (Quezon Province): Bulalon Cave, Burdeos (v.2010 JMA Encinares, UPLBMNH BLA-00441-00442, nymphs); Cave 3-4 (v.2011 JMA Encinares, UPLBMNH BLA-00532, nymph).
Front femur A2, pulvilli on 4 proximal tarsomeres, tarsal claw simple and symmetric, arolium present. Quite similar to nymphs of H. silbergliedi except for the broad lateral white margin of the pronotum and completely black abdomen.
Accidentals. This species has been previously recorded only from Bulalon Cave (
The authors would like to express their gratitude to the UPLB-MNH staff for the allowing the first author to tag along during their expeditions and for the use of their laboratory equipment and space during the examination of the specimens; Dr. Aimee Lynn B. Dupo for the use of camera that generated high quality images; IBS Cave Ecology Laboratory, also for the use of space and facilities; Mr. John Mark A. Encinares for the information regarding the specimens from Bulalon and Mapanghe caves; Dr. George W. Beccaloni, Dr. Darren J. Mann and Dr. Christine A. Nalepa for sending references otherwise unavailable to us; and lastly, the local government of Burdeos, Quezon for the permission to conduct the study and the Polillo Ecology Wardens for their support during the fieldwork, without which these expeditions will not be possible. Funding for initial fieldwork in Polillo was received from the UPLB Basic Research Project (No. 88-D66-00) of the second author.