ZooKeys 822: 17-31 (20 | 9) A peer-rev iewed open-access journa I

doi: 10.3897/zookeys.822.30927 RESEARCH ARTICLE #ZooKey

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Three new species of the spider genus Luzonacera Li &
Li, 2017 from Philippines (Araneae, Psilodercidae)

Wan-Jin Chang'’, Fengyuan Li', Shuqiang Li!

| Institute of Zoology, Chinese Academy of Sciences, Beijing 100101, China 2 University of Chinese Academy
of Sciences, Beijing, China

Corresponding author: Shugiang Li (lisq@ioz.ac.cn)

Academic editor: Y. Marusik | Received 30 October 2018 | Accepted 15 January 2019 | Published 4 February 2019
http://zoobank.org/767 C4AC0-87 B4-4E 10-AA64-523D8DFFD87D

Citation: Chang W-J, Li F, Li S (2019) Three new species of the spider genus Luzonacera Li & Li, 2017 from
Philippines (Araneae, Psilodercidae). ZooKeys 822: 17-32. https://doi.org/10.3897/zookeys.822.30927

Abstract

Three new species of Luzonacera Li & Li, 2017 are described: L. francescoballarini Li & Li, sp. n. (49),
L. lattuensis Li & Li, sp. n. (SQ) and L. peterjaegeri Li & Li, sp. n. (4Q). Prior to this study, the genus
was known by two species, both from Luzon Island, Philippines. So far, the genus and all five species are

endemic to Luzon Island and can be found in dry or humid caves in a dark environment.

Keywords

cave, endemic, Luzon Island, Southeast Asia, tropical

Introduction

The spider family Psilodercidae Machado, 1951 contains eleven genera and 116 species
(World Spider Catalog 2018, Li and Quan 2017). All species are restricted to tropical Asia
and known from Sri Lanka and India to Philippines (World Spider Catalog 2018). Cur-
rently, five species of Psilodercidae belonging to four genera are known to occur in Philip-
pines (World Spider Catalog 2018): Psiloderces egeria Simon, 1892 from Luzon, Althepus
noonadanae Brignoli, 1973 from Mindanao, Leclercera negros Deeleman-Reinhold, 1995
from Negros, and Luzonacera chang Li & Li, 2017 and L. duan Li & Li, 2017 from Luzon.

The recently described genus Luzonacera Li & Li, 2017 was known from two species,
L. chang Li & Li, 2017 and L. duan Li & Li, 2017 (World Spider Catalog 2018). While
studying new material collected on Luzon Island, we recognized three new species of
the genus. The goal of this paper is to provide detailed descriptions of these new species.

Copyright Wan-Jin Chang et al. 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.

18 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

Materials and methods

All specimens were collected in Luzon Island and preserved in 95% ethanol solution. All
types are deposited in the Institute of Zoology, Chinese Academy of Sciences in Beijing
(IZCAS) and Senckenberg Research Institute in Frankfurt (SMF). A Leica M205 C ster-
eomicroscope was used to measure and examine the specimens. Morphological details of
the specimens were studied with an Olympus BX41 compound microscope. An Olympus
C7070 wide zoom digital camera (7.1 megapixels) mounted on an Olympus SZX12 ster-
eomicroscope was used to take photos. The images were generated using Helicon Focus
6.7.1 image stacking software and further revised with Adobe Photoshop. Leg measure-
ments are shown as total length (femur, patella, tibia, metatarsus, and tarsus). Leg segments
were measured from their retrolateral side except for L. peterjaegeri sp. n. which was meas-
ured from the prolateral side. All measurements are given in millimetres (mm). Terminol-
ogy follows that of Li et al. (2014), Tong and Li (2007) and Deeleman-Reinhold (1995).
The extraction of genomic DNA from legs followed Li and Li (2018). Primer
sets for the PCR and cycle sequencing reactions used for cytochrome c oxidase subu-
nit I (COI) in this study are from Folmer et al. (1994). All sequences were analysed
using BLAST. The GenBank accession numbers are provided in Table 1. The COI
dataset of the three sequences obtained in this study and two sequences from Gen-
Bank were aligned using MAFFT version 7 (http://mafft.cbrc.jp/ alignment/server/).
MEGA7.0.16 (Kumar et al. 2016) was used for subsequent manual adjustment of the
sequences and calculation of pairwise comparisons of uncorrected K2P-distances.

Table |. The accession numbers for each species in this paper.

Species Length (bp) GenBank accession number
Luzonacera francescoballarini sp. n. 651 MK238752
Luzonacera lattuensis sp. n. 651 MK238753
Luzonacera peterjaegeri sp. n. 651 MK238754
Taxonomy

Family Psilodercidae Machado, 1951

Genus Luzonacera Li & Li, 2017

Type species. Luzonacera chang Li & Li, 2017

Emended diagnosis. Luzonacera resembles Althepus Thorell, 1898 and Leclercera
Deeleman-Reinhold, 1995. However, Luzonacera can be differentiated by the combi-
nation of the following characteristics: 1) absence of a conductor (versus presence of a
conductor in both Althepus and Leclercera); 2) absence of a retrolateral protrusion on the
tibia or cymbium of the male palp (versus presence of a retrolateral protrusion on the tib-
ia or cymbium of the male palp in Althepus and Leclercera); 3) remarkably inflated tibia
of the male palp; 4) pyriform bulb with spirally extended embolus; and 5) two pairs of
spermathecae, the lateral spermathecae with longer stalks than the medial spermathecae.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 19

Composition. L. chang Li & Li, 2017 (the type species), L. duan Li & Li, 2017,
L. francescoballarini sp. n., L. lattuensis sp. n. and L. peterjaegeri sp. n.
Distribution. Philippines.

Illustrated key to the males of Luzonacera

1 Embolus and bulb equal in length; embolus and bulb ratio approximately

SGU AMtO MET MI) so wata nwo teveodnrnr stead ooraoscveey baceesnasiauete tinouniitev enn L. chang
= Embolus short (bulb ca. 2 times longer than the embolus); embolus and bulb

Fanos 1A 005.7 Obie NSW panarser ct diauneedt ats olseteau tates ob odelse Ere collate ueusaa tt 2
2 Slichticonstriction-of thecentraliparwot bulbA Pig Alii) ss cs.sccpsles csapavdespanndss
SSR Aaa sada Sea hae oh ates Vases acs aoeawtahn eee L. francescoballarini sp. n.
Pronounced constriction of the central part of bulb (Fig. Lii, iv, v) ......... 3
Bulb with smooth surface dorsally (Fig. lv) ww... L. peterjaegeri sp. n.
= Bulbswithanoreh NO) dorsally, (Pig wl O2) 1) 20. sccvedennidecoerearcsvaninsrateess 4
The tip of the bulb without protrusion (PT) (Fig. ii)
The tip of the bulb with protrusion (PT) (Fig. liv) .........

ww |

aX

Figure 1. Prolateral view of male palp bulbs of Luzonacera species: i L. chang ii L. duan iii L. francescob-
allarini sp. n. in L. lattuensis sp. n. and v L. peterjaegeri sp. n. Abbreviations: PT = protrusion, NO = notch.

Illustrated key to the females of Luzonacera

1 Two paitsiof-similarspermathecaes@=d) =. tsctelent iss cere et eee 2
7 Two pairs of dissimilar spermathecae (medial pair oblique, tube-shaped) (e)...
wns dos arsngabia bn tash's eds enspiny Wetaascnnawell ete easasieW ec adadenhte Meenas aaeenne L. peterjaegeri sp. n.

20 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

2 Spermathecae without globose distal part (with swollen distal ends) (Fig. 2a)
Oe LO eR Sent rea eae ee oe OER OR DeLee Ser L. chang
— Spermathecae with globose distal part (Fig. 2b-d) oo... eeeseeseceeeeeeeeees 3
3 Two pairs of spermathecae pointed almost the same direction (Fig. 2b, d) ..4
~ Two pairs of spermathecae pointed opposite directions (Fig. 2c) 0... eee
ee Re ee oy te oan eon L. francescoballarini sp. n.
=: Relatively short stalks of medial spermathecae; distal part and stalk ratio ap-
Pro <imncite ly OR sei DA): caries. worse. sites eames eae ea L. lattuensis sp. n.
7 Relatively long stalks of medial spermathecae; distal part and stalk ratio ap-
prominiately O82 (Rice 2) tenes. m5... 5 0m... amma nes asa heme ser mn, L. duan

Figure 2. Female internal genitalia of five different species of Luzonacera: a L. chang b L. duan € L.
francescoballarini sp. n. d L. lattuensis sp. n. and e L. peterjaegeri sp. n. Abbreviation: SP = spermathecae.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 21

Luzonacera francescoballarini Li & Li, sp. n.
http://zoobank.org/EEE59EF0-7 FF6-4F7 D-BB5E-GFE76729FB76
Figs 3, 4, 9, 10

Types. Holotype: ¢ (IZCAS), Philippines, Luzon Island, Bulacan Province, San
Miguel City, near Biak-Na-Bato National Park, Bayukbok Cave, 15°10'5.4"N,
121°5'4.3"E, 125 m, 21.V.2015, E Ballarin and Y. Li. Paratypes: 14, 19 (IZCAS),
same data as holotype.

Etymology. The species is named after Francesco Ballarin, who collected the type
series; name in genitive case.

Diagnosis. Luzonacera francescoballarini sp. n. resembles L. lattuensis sp. n. in hav-
ing a short embolus, and two pairs of twisted spermathecae globose at distal parts.
Males can be distinguished from the latter species by the smooth dorsal surface of the
bulb (Figure 3B); females can be distinguished by having longer spermathecae (Figure
4A; versus shorter in L. /attuensis sp. n. in Figure GA).

Description. Male (Holotype). Total length 4.81; carapace 1.60 long, 1.28 wide;
abdomen 3.20 long, 0.96 wide. Colour faded. Carapace round, pale yellow, with ovoid
brown patch medially and rounded brown patch posterior to ocular area. Fovea shal-
low. Anterior margin of thoracic region distinctly elevated. Chelicerae light brown
with lamina, promargin with a single tooth and retromargin with two small teeth
(Figure 7A). Clypeus slanting, light brown. Labium slanting, pale brown. Sternum
pale brown with large patch of brown spots medially. Abdomen elongated with com-
plex patterns dorsally and ventrally. Legs light brown; measurements: I 22.44 (6.73,
0:64, 6741'523.21; 5:45), Ih 17-6345.13, 0:64; 5.13;,5.45; 1.28); 1 12:05-(3.53).0.32,
3.40, 3.20, 1.60), IV 23.07 (7.05, 0.64, 7.05, 6.41, 1.92). Palp (Figure 3A—E): tibia
swollen at the base, length/width = 2.40; cymbium swollen with distal protrusion and
numerous long setae; length/width = 2.33; bulb light brown, pyriform; embolus forms
a slender spiral extending from tip of bulb.

Female (paratype). Similar to male in coloration and general features but slightly
larger (Figure 4D, E). Measurements: total length 3.59; carapace 1.20 long, 1.40
wide; abdomen 2.20 long, 0.80 wide. Leg measurements: I 14.11(4.17, 0.32, 4.17,
ANT; 1.28), 10 10-883 32505,0.92,79:95;-3-21,.0-32) 2 9 26 (56,050,200 82.60;
1.00), IV 14.06 (4.49, 0.40, 4.17, 4.00, 1.00). Internal genitalia: two pairs of slender
spermathecae with long stalks (ca. 6 times longer than distal globular parts), sper-
mathecae distal parts not wider than basal width of stalks, both pairs equal in width
(Figure 4A).

Distribution. Type locality only (Figure 10).

Natural history. Collected in a dark and rather humid cave, close to the ground,
along the wall of the cave with huge rocks.

Comments. Based on the 651 bp aligned sequences, the COI uncorrected K2P-
distance between L. francescoballarini sp. n. and L. chang is 13.5%, between L. francesco-
ballarini sp. n. and L. duan is 15.0%, between L. francescoballarini sp. n. and L. lattuensis
sp. n. is 14.9%, and between L. francescoballarini sp. n. and L. peterjaegeri sp. n. is 13.9%.

22 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

Figure 3. Luzonacera francescoballarini sp. n., male holotype A palp bulb, retrolateral view B palp bulb,
ventral view C palp bulb, prolateral view D palp, prolateral view E palp, retrolateral view. Abbreviations:

EM = embolus; CP = cymbial protrusion.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 23

Figure 4. Luzonacera francescoballarini sp. n., male holotype and female paratype A internal genitalia,

dorsal view B female epigastric furrow, ventral view C male habitus, dorsal view D female habitus, dorsal

view E female habitus, ventral view. Abbreviation: SP = spermathecae.

24 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

Luzonacera lattuensis Li & Li, sp. n.
http://zoobank.org/0D355C5B-450C-4F82-805E-0062E14381F0
Figs:5,°6, 9,10

Types. Holotype: 3 (IZCAS), Philippines, Luzon Island, Cagayon Province, Tugue-
garao City, Penablanca Village, Lattu-Lattuc Cave, 17°42'23"N, 121°49'2"E, 111 m,
31.V.2015, EF Ballarin and Y. Li. Paratypes: 14), 12 (IZCAS), same data as holotype.

Etymology. ‘The species name is an adjective referring to the type locality.

Diagnosis. Both sexes of L. /attuensis sp. n. and L. francescoballarini sp. n. are very
similar. Males of L. lattuensis sp. n. can be distinguished from L. francescoballarini sp.
n. by the bulb with a dorsal notch (Figure 5A) and a relatively longer cymbium tip
(Figure 5E); females can be distinguished by having shorter spermathecae with more
widely spaced bases (Figure 6A; versus longer spermathecae with more narrowly spaced
bases in L. francescoballarini sp. n. in Figure 7A).

Description. Male (Holotype). Total length 3.85; carapace 1.28 long, 0.96 wide;
abdomen 2.56 long, 0.75 wide. Carapace round and brown, with three longitudinal
brown bands; the central band is 3 times wider than the lateral bands (Figure 6C).
Fovea shallow, brown. Anterior margin of thoracic region distinctly elevated. Cheli-
cerae brown with lamina, promargin with one tooth and retromargin with two small
teeth (Figure 9B). Clypeus slanting, brown with two pale rounded areas laterally and
two triangular projections basally. Labium slanting, dark brown. Sternum pale brown
with three dark brown patches laterally. Abdomen elongated, with complex patterns
dorsally and ventrally. Legs brown with white annulations; measurements: I & II
missing, III 7.80 (2.24, 0.32, 2.24, 2.00, 1.00), IV missing. Palp (Figure 5A—E): tibia
swollen at the base, length/width = 2.50; cymbium with distal protrusion, length/
width = 2.0; bulb light brown, pyriform; embolus forms a slender spiral extending
subapically from bulb.

Female (paratype). General features and coloration are similar to the male, but
the female is slightly larger (Figure 6D, E). Measurements: total length 3.81; carapace
1.25 long, 1.00 wide; abdomen 2.56 long, 1.40 wide. Leg measurements: I 11.05
(3225; 0,40): 3320332204 1200), 10 10:57 G20;0.32, 2:83).391 ,:0:96), Ih 674-200;
0.32, 1.92, 1.75, 0.75), IV 13.35 (3.75, 0.31, 3.80, 4.49, 1.00). Internal genitalia: two
pairs of twisted spermathecae, medial spermathecae with globose distal parts and short
stalks, and lateral spermathecae with globose distal parts and long stalks (stalks ca. 3
times longer than distal parts) (Figure 6A).

Distribution. Type locality only (Figure 10).

Natural history. Collected close to the ground along the wall of a dark, rather dry
and dusty secondary cave with huge rocks.

Comments. Based on the 651 bp- aligned sequences, the COI uncorrected K2P-
distance between L. /attuensis sp. n. and L. chang is 12.4%, between L. lattuensis sp. n.
and L. duan is 11.5%, and between L. /attuensis sp. n. and L. peterjaegeri sp. n. is 13.6%.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 25

Figure 5. Luzonacera lattuensis sp. n., male holotype A palp bulb, retrolateral view B palp bulb, ventral
view C palp bulb, prolateral view D palp, prolateral view E palp, retrolateral view. Abbreviations: EM =
embolus; CP = cymbial protrusion; NO = notch.

26 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

PEN IRR

Figure 6. Luzonacera lattuensis sp. n., male holotype and female paratype A internal genitalia, dorsal
view B female epigastric furrow, ventral view C male habitus, dorsal view D female habitus, dorsal view

E female habitus, ventral view. Abbreviation: SP = spermathecae.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 27

Figure 7. Luzonacera peterjaegeri sp. n., male holotype A right palp bulb, retrolateral view B right palp
bulb, ventral view C right palp bulb, prolateral view D right palp, prolateral view E right palp, retrolateral

view. Abbreviations: EM = embolus; CP = cymbial protrusion.

Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

28

0.01mm

0.5mm

0.5mm

Figure 8. Luzonacera peterjaegeri sp. n., male holotype and female paratype A internal genitalia, dorsal
view f female epigastric furrow, ventral view C male habitus, dorsal view D female habitus, dorsal view

E female habitus, ventral view. Abbreviation: SP = spermathecae.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... 29

Luzonacera peterjaegeri Li & Li, sp. n.
http://zoobank.org/ B57BC1C9-4645-4662-90C1-CE3FFC211E96
Figs 7-9

Types. Holotype: 3 (SME), Philippines, Northern Luzon Island, Teresita State, Ca-
gayan Province, Lower Kimmabalyu Cave, 18°11'35.4"N, 121°52'10.3"E, 22.1.2015,
H. Steiner. Paratypes: 14, 29 (SMF), same data as holotype.

Etymology. ‘The species is named after Peter Jager in honour of his contribution to
the study of spiders from Asia; name in genitive case.

Diagnosis. Luzonacera peterjaegeri sp. n. can be distinguished from all other known
species of the genus by a distinct constriction on the central part of the bulb (Figure
7A); females can be distinguished by two types of spermathecae: one pair of slender
spermathecae bearing a globose distal part, and one pair of oblique, tube-shaped sper-
mathecae (Figure 8A; versus both pairs of spermathecae bearing a globose distal part
in other species). Moreover, both sexes of L. peterjaegeri sp. n. have longer chelicerae
(Figure 8; versus shorter chelicerae in other species).

Description. Male (Holotype). Total length 4.00; carapace 1.50 long, 1.40 wide;
abdomen 2.50 long, 1.00 wide. Carapace round, pale brown, with three longitudinal
brown bands, with the middle band 3 times wider than the lateral bands (Figure 8C).
Fovea shallow and brown. Anterior margin of thoracic region distinctly elevated.
Chelicerae long, brown with lamina, promargin with one tooth, and retromargin
with two small teeth (Figure 9C). Clypeus brown with two pale rounded areas later-
ally and two relatively longer protrusions basally. Labium brown. Sternum brown
with three dark brown patches laterally. Abdomen elongated, with complex patterns

Figure 9. Cheliceral retromargin A L. francescoballarini sp. n. B L. lattuensis sp. n. C L. peterjaegeri sp. n.

Abbreviations: PT = promargin teeth; RT = retromargin teeth; CL = cheliceral lamina.

30 Wan-Jin Chang et al. / ZooKeys 822: 17-31 (2019)

dorsally and ventrally. Leg measurements: all legs missing. Right palp (Figure 7A—
E): tibia swollen at the base, length/width = 2.25; cymbium with distal protrusion,
length/width = 3.4; bulb light brown, pyriform; embolus forms a slender spiral elon-
gating terminally from bulb.

Female (paratype). General features and coloration are similar to male, but the
female is slightly larger (Figure 8D, E). Measurements: total length 4.17; carapace
1.60 long, 1.28 wide; abdomen 2.56 long, 1.5 wide. Legs missing. Internal genitalia:
two pairs of spermathecae, one pair of spermathecae globose distally with long stalks
(ca. 6 times longer than distal parts), the other pair are oblique, tube-shaped sper-
mathecae (Figure 8A).

Distribution. Type locality only (Figure 10).

Comments. Based on the 651 bp aligned sequences, the COI uncorrected K2P-
distance between L. peterjaegeri sp. n. and L. chang is 15.9%, and between L. peterjae-
geri sp. n. and L. duan is 13.9%.

115°E 120°E 125°E 130°E
20°N

200 100 0 200 KM
ee

15°N

10°N

Figure 10. Distribution of five Luzonacera in Philippines | Luzonacera francescoballarini sp. n. 2 L. lat-
tuensis sp. n. 3 L. peterjaegeri sp. n. 4 L. duan § L. chang.

Three new species of the spider genus Luzonacera Li & Li, 2017 from... Je

Acknowledgements

The manuscript benefited greatly from comments by Drs Yuri Marusik (Magadan,
Russia), Mark Harvey (Western Australia, Australia), and Yanfeng Tong (Shenyang,
China). Sarah Crews (San Francisco, USA) kindly checked the English. We are in-
debted to Francesco Ballarin (Tokyo, Japan), Helmut Steiner (Hanau, Germany), and
Peter Jaeger (SMF) who provided the specimens examined in this study. This study was
supported by the National Natural Science Foundation of China (NSFC-31530067)
to Shugiang Li.

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