High level of endemism in Haiti’s last remaining forests: a revision of Modisimus (Araneae: Pholcidae) on Hispaniola, using morphology and molecules
Author
Huber, Bernhard A.
Author
Fischer, Nadine
Author
Astrin, Jonas J.
text
Zoological Journal of the Linnean Society
2010
2010-01-25
158
2
244
299
http://dx.doi.org/10.1111/j.1096-3642.2009.00559.x
journal article
10.1111/j.1096-3642.2009.00559.x
0024-4082
5438272
MODISIMUS FEMORATUS
BRYANT, 1948
(
Figs 17, 18
,
38
,
63
,
83
,
94
,
104–107
,
124–126
,
195
)
Modisimus femoratus
Bryant, 1948: 367–368
, figs 39, 42.
M. glaucus
Simon
:
Bryant, 1948: 369–370
, figs 38, 41, 56; misidentification? (See note below.)
Note:
Two males from
Puerto Plata
identified by
Bryant (1948)
as
M. glaucus
Simon
are here tentatively assigned to
M. femoratus
Bryant. However
, they might indeed be
M. glaucus
; equally they might neither be
M. glaucus
nor
M. femoratus
, or
M. femoratus
could in fact be a junior synonym of
M. glaucus
(also see notes on
type
species above).
Type:
Male
holotype
from
Sánchez
[~
19°13.8
′
N
,
69°36.8
′
W
],
Samaná Prov.
,
Dominican Republic
; with labels ‘Dom. Rep.;
Sánchez
,
July 1938
,
Darlington’
and ‘
Modisimus femoratus
♂
–
holotype
B.0023 Bryant’, in
MCZ
, examined
.
Diagnosis:
Medium-sized species with slightly elongated abdomen (
Figs 17, 18
), male chelicerae with pair of small projections, with weakly modified hairs (
Figs 83
,
125
; similar to
M. jima
sp. nov.
and
M. angulatus
sp. nov.
), and distinctive epigynum (
Fig. 38
; similar to
M. angulatus
sp. nov.
). Distinguished from
M. angulatus
sp. nov.
by pattern on sternum (lateral border not broken), dorsally evenly curved abdomen, and male cheliceral projections slightly closer together; distinguished from
M. jima
sp. nov.
by longer abdomen, indistinct ventral procursus notch, and slightly different epigynum.
Male (near Sánchez):
Total length, 2.6; carapace width, 1.1. Leg 1: 29.4 (7.8 + 0.4 + 7.3 + 12.0 + 1.9); tibia 2, 4.8; tibia 3, 4.0; tibia 4, 4.4. Tibia 1 L/d: 63. Habitus as in
Figures 17 and 18
: carapace ochreyellow, ocular area posteriorly brown, clypeus laterally light brown; sternum with two brown longitudinal bands, medially whitish, lateral margins ochre-yellow; legs light brown, tips of femora and tibiae whitish; abdomen bluish grey, with dark spots dorsally and laterally; large light-brown genital area, dark brown in front of spinnerets, with blue mark between genital area and spinnerets. Ocular area strongly elevated; thoracic furrow distinct.
PME
–
PME
, 80 Mm;
PME
diameter, 105 Mm;
PME
–
ALE
, 115 Mm;
AME
very tiny (
Fig. 83
), lenses not clearly visible in dissecting microscope. Sternum wider than long (0.7/0.5), unmodified. Chelicerae with indistinct round humps frontally set with a few basally thickened hairs (
Figs 83
,
125
). Palps as in
Figure 124
, coxa with light retrolateral apophysis, femur with rounded proximal and pointed distal ventral apophyses, procursus with dark dorsal spine-like process (see also
Fig. 94
) and membranous structures distally, and bulb with large, weakly curved apophysis. Legs with spines in two ventral rows on femora 1 and 2 (~45–55 spines in each row on femur 1; 20–25 spines in each row on femur 2), some spines on elevated bases, retrolateral trichobothrium on tibia 1 at 8%; all femora with higher than usual density of short vertical hairs, but not very dense; curved hairs dorsally on tibiae and metatarsi 1–3; prolateral trichobothrium missing on tibia 1, but present on all other tibiae. Tarsus 1 with ~30 pseudosegments.
Variation:
Tibia
1 in
53 other males: 5.8–8.8 (mean 7.3). Small males tend to have fewer and smaller spines on their femora; some males have curved hairs on tibiae only; some males also have white spots on their abdomen. The
type
is in poor condition, with legs fragmented, both palps and abdomen detached, and with legs 1 missing. Males from Loma Quita Espuela and
Puerto Plata
appear identical, but are smaller (tibia
1 in
ten males
, 5.0–6.4; mean, 5.5; cf. females from that locality); they are therefore only assigned tentatively. The material from
Dajabón Prov.
is assigned tentatively because the male cheliceral prominences are slightly more distal, the procursus tip differs slightly, and they are smaller (tibia 1: 5.1, 5.7).
Female:
In general similar to male, but with more distinctive brown lateral marks on clypeus. Tibia
1 in
37 females
: 3.6–5.4 (mean 4.7). Epigynum, roughly rectangular, with posterior light area flanked by lateral sclerites (
Fig. 38
); frontally, a triangular sclerite; dorsal view as in
Figures 63
and
126
, with a prominent frontal membranous structure. Females from Loma Quita Espuela and
Puerto Plata
tend to be smaller: tibia
1 in
12 females
: 3.5–4.3 (mean 3.95); in those from Loma Quita Espuela, the epigynum is slightly longer than usual. Females from
Dajabón Prov.
have extremely similar epigyna; tibia 1, 3.8 and 4.0.
Distribution and habitat:
This species is widely distributed in the north-eastern
Dominican Republic
(
Fig. 195
). It was sometimes found to share its habitat with
M. kiskeya
sp. nov.
(among vegetation), but lives closer to the ground, and also among rocks and under dead leaves.
Material examined:
Dominican Republic
:
Samaná Prov.
,
Sánchez
,
1♂
,
holotype
above
;
near
Sánchez
, degraded forest at road to
Las Terrenas
(
19°14.5
′
N
,
69°35.9
′
W
), domed webs in vegetation,
290 m
a.s.l.
,
11 November 2005
(
B.A. Huber
),
7♂
and
9♀
(
ZFMK
,
DR 30
)
;
same data,
1♀
, in pure ethanol (
ZFMK
,
DR 100-30
)
;
degraded forest near
Saltos de Limón
(~
19°16.6
′
N
,
69°26.5
′
W
), ~
120 m
a.s.l.
, in low vegetation,
11 November 2005
(
B.A. Huber
),
8♂
,
7♀
and
two juveniles
(
ZFMK
,
DR 33
)
;
same data,
1♂
,
5♀
and
one juvenile
, in pure ethanol (
ZFMK
,
DR 100-27
)
;
south of
Las Gelaras
(
19°12.8
′
N
,
69°13.1
′
W
), forest above rocks, low vegetation, near ground,
20 m
a.s.l.
,
12 November 2005
(
B.A. Huber
),
11♂
and
7♀
(
ZFMK
,
DR 36
)
;
same locality,
1♂
, in pure ethanol, together with
one juvenile
of an unidentified species (
Modisimus
sp. 3
;
Table 4
) (
ZFMK
,
DR 100-33
)
.
María Trinidad Sánchez Prov.
, near
La Entrada
, forest above rocks at
Santuario de La Virgen
(
19°34.9
′
N
,
69°54.0
′
W
),
15 m
a.s.l.
,
12 November 2005
(
B.A. Huber
),
6♂
,
3♀
and
one juvenile
(
ZFMK
,
DR 41
)
.
Monte Plata Prov.
, near
Yamasá
(
18°45.9
′
N
,
70°01.2
′
W
), degraded forest along river,
70 m
a.s.l.
, low vegetation,
20 November 2005
(
B.A. Huber
),
14♂
,
3♀
and
four juveniles
(
ZFMK
,
DR 84
/85)
;
same data,
1♀
, in pure ethanol (
ZFMK
,
DR 100-20
)
; near
Monte Plata
(
18°48.7
′
N
,
69°47.1
′
W
), degraded forest along river,
60 m
a.s.l.
, near ground,
21 November 2005
(
B.A. Huber
),
4♂
and
1♀
(
ZFMK
,
DR 91
)
.
Hato Mayor Prov.
, north of
Manchado
(
18°50.2
′
N
,
69°18.5
′
W
), degraded forest near river,
150 m
a.s.l.
, vegetation,
21 November 2005
(
B.A. Huber
),
2♂
,
5♀
and
four juveniles
(
ZFMK
,
DR 94
)
; forest between
Hato Mayor
and
Sabana de la Mar
,
2 July 1935
(
W. G. Hassler
),
2 ♀
in poor condition (
AMNH
)
.
El Seíbo Prov.
, near
Miches
(
18°56.8
′
N
,
69°05.2
′
W
), forest with plantations, ~
300 m
a.s.l.
, low vegetation,
22 November 2005
(
B.A. Huber
),
5♂
,
5♀
and
three juveniles
(
ZFMK
,
DR 99
)
.
Figures 17–37.
Habitus photos of males (except 33), showing typical and distinctive characters. 17, 18.
M. femoratus
. 19.
M. jima
sp. nov
.
20.
M. kiskeya
sp. nov.
21, 22.
M. angulatus
sp. nov.
23, 24.
M. epepye
sp. nov.
25.
M. mango
sp. nov.
26.
M. cienaga
sp. nov.
27–29.
M. seguin
sp. nov.
(arrows point to callus-like area). 30–34.
M. vittatus
. 35.
M. palvet
sp. nov.
36.
M. berac
sp. nov.
37.
M. paraiso
sp. nov.
Figures 38–62.
Epigyna in ventral view. 38.
M. femoratus
(arrow points to frontal sclerite). 39.
M. jima
sp. nov
.
40.
M. angulatus
sp. nov.
41.
M. kiskeya
sp. nov.
42.
M. toma
sp. nov
.
43.
M. vittatus
(arrow points to distinctive posterior sclerite). 44.
M. makandal
sp. nov.
45.
M. fuscus
. 46.
M. epepye
sp. nov.
47.
M. seguin
sp. nov.
48.
M. roumaini
sp. nov.
49, 50.
M. enriquillo
sp. nov.
from near Polo (49) and from south of Barahona (50) (arrows point to sternum apophyses). 51, 52.
M. tiburon
sp. nov.
53.
M. leprete
sp. nov.
54.
M. macaya
sp. nov.
55.
M. bachata
sp. nov.
56.
M. cuadro
sp. nov.
57.
M. pelejil
sp. nov.
58.
M. mango
sp. nov.
59.
M. berac
sp. nov.
60.
M. palvet
sp. nov.
61.
M. miri
sp. nov.
62.
M. paraiso
sp. nov.
Figures 63–82.
Cleared and stained epigyna in dorsal view. 63.
M. femoratus
. 64.
M. jima
sp. nov
.
65.
M. toma
sp. nov
.
66.
M. vittatus
. 67.
M. makandal
sp. nov.
68.
M. fuscus
. 69.
M. epepye
sp. nov.
70.
M. seguin
sp. nov.
71.
M. roumaini
sp. nov.
72.
M. enriquillo
sp. nov.
73.
M. tiburon
sp. nov.
74.
M. leprete
sp. nov.
75.
M. macaya
sp. nov.
76.
M. mariposas
sp. nov.
77.
M. bachata
sp. nov.
78.
M. cuadro
sp. nov.
79.
M. pelejil
sp. nov.
80.
M. mango
sp. nov.
81.
M. berac
sp. nov.
82.
M. miri
sp. nov.
Figures 83–93.
SEM images of selected characters. 83–85. Male prosomata in frontal view:
M. femoratus
(83; arrow points to tiny anterior median eye, AME),
M. toma
sp. nov.
(84), and
M. cienaga
sp. nov.
(85). 86. Male prosoma, dorsal view (
M. kiskeya
sp. nov.
). 87. Ocular area and carapace, frontodorsal view (
M. cienaga
sp. nov.
). 88–91. Male cheliceral hairs:
M. seguin
sp. nov.
(88),
M. palvet
sp. nov.
(89), and
M. toma
sp. nov.
(90, 91). 92, 93. Male gonopore,
M. cienaga
sp. nov.
(92),
M. palvet
sp. nov.
(93). Scale bars: 20 Mm (89); 40 Mm (90, 92, 93); 60 Mm (88, 91); 100 Mm (85, 87); 300 Mm (83, 84, 86).
Figures 94–103.
SEM images of selected characters. 94. Left procursus, retrolateral view (
M. femoratus
). 95. Right procursus and distal bulbal elements, retrolaterodorsal view (
M. kiskeya
sp. nov.
). 96. Left palp, retrolateral view (
M. toma
sp. nov.
). 97 Right procursus and distal bulbal elements, distal view (
M. toma
sp. nov.
). 98. Left procursus, retrolateral view (
M. seguin
sp. nov.
). 99. Left procursus, retrolateral view (
M. cuadro
sp. nov.
). 100. Right procursus and distal bulbal elements, distal view (
M. cienaga
sp. nov.
). 101. Palpal tarsal organ (
M. cienaga
sp. nov.
). 102. Bulb with putative sperm duct opening (
M. cienaga
sp. nov.
). 103. Left procursus tip, retrolateral view (
M. cienaga
sp. nov.
). Abbreviations: b, genital bulb; ba, bulbal apophysis; p, procursus. Scale bars: 8 Mm (101); 30 Mm (100, 102, 103); 60 Mm (94, 95, 97); 100 Mm (96, 98, 99).
Figures 104–114.
SEM images of selected characters. 104–107. Epigynum of
M. femoratus
, ventral view, without plug (104), ventroposterior view, with plug (105), and ventral views, with plug (106, 107). 108–110. Epigynum of
M. toma
sp. nov.
, ventral view without plug (108), ventral view with plug (109), and ventroposterior view with plug (110). 111, 112. Epigynum of
M. cuadro
sp. nov.
, ventral (111) and ventroposterior (112) views. 113, 114. Epigyna of
M. cienaga
sp. nov.
(113) and
M. palvet
sp. nov.
(114), ventral views. Abbreviation: gp, ‘genital plug’. Scale bars: 100 Mm (104, 105, 107–114); 200 Mm (106).
Assigned tentatively:
Dajabón Prov.
, near El Pino (
19°24.7
′
N
,
71°29.9
′
W
), degraded forest along river,
160 m
a.s.l.,
15 November 2005
(B.A. Huber),
1♂
and
2♀
(
ZFMK
, DR 54); south of Restauración (
19°16.8
′
N
,
71°42.1
′
W
), degraded forest along brook,
630 m
a.s.l.,
16 November 2005
(B.A. Huber),
1♂
and
one juvenile
(
ZFMK
, DR 57).
Puerto Plata Prov.
: south of
Puerto Plata
, forest along path towards Isabel Torres National Park (
19°46.4
′
N
,
70°42.3
′
W
),
350 m
a.s.l.,
14 November 2005
(B.A. Huber),
8♂
and
6♀
(
ZFMK
, DR 49); same data,
1♂
and
5♀
, in pure ethanol, one of them sequenced and found to be considerably distant from
M. femoratus
(see
Modisimus
sp.
2
in
Table 4
) (
ZFMK
, DR 100–39);
Puerto Plata
,
July–August 1941
(Hurst),
2♂
(
MCZ
26.669).
Duarte Prov.
, Reserva Científica Loma Quita Espuela at ~
500 m
a.s.l. (
19°21.5
′
N
,
70°09
′
W
),
10 November 2005
(B.A. Huber),
2♂
and
6♀
(
ZFMK
, DR 25); same data,
4♀
, in pure ethanol, one of them sequenced and found to be considerably distant from
M. femoratus
(see
Modisimus
sp.
1
in
Table 4
) (
ZFMK
, DR 100-23). ‘
75 km
N Santo Domingo’,
17 July 1935
(Hassler),
2♂
and
one juvenile
(
AMNH
).