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+
+
+
+A second endemic land mammal for the Hawaiian Islands: a new genus and species of fossil bat (Chiroptera: Vespertilionidae)
+
+
+
+Author
+
+Ziegler, Alan C.
+Francis G. Howarth, & Department of Natural Sciences, Bishop Museum, Honolulu, Hawai‘i. & Francis G. Howarth, & Deceased.
+
+
+
+Author
+
+Howarth, Francis G.
+Francis G. Howarth, & Department of Natural Sciences, Bishop Museum, Honolulu, Hawai‘i.
+
+
+
+Author
+
+Simmons, Nancy B.
+Department of Mammalogy, Division of Vertebrate Zoology, American Museum of Natural History, New York.
+
+text
+
+
+American Museum Novitates
+
+
+2016
+
+2016-03-21
+
+
+2016
+
+
+3854
+
+
+1
+52
+
+
+
+
+http://www.bioone.org/doi/10.1206/3854.1
+
+journal article
+10.1206/3854.1
+0003-0082
+5368539
+
+
+
+
+
+Synemporion keana
+,
+
+gen. and sp. nov.
+
+
+
+Lava-tube Bat
+
+Figures 2–8
+, tables 1–3
+
+
+
+
+
+HOLOTYPE
+:
+BPBM 159269
+, a nearly complete skeleton of an
+adult
+individual including skull, both dentaries, and partial body skeleton with both scapulae, humeri, ulnae, radii, and femora; right tibia and partial innominate with 2 fused sacral vertebrae; 8 metacarpals; 2 manual phalanges; 1 pedal phalanx; atlas; 2 thoracic vertebrae; and 1 rib. This individual was collected on
+
+16 September 1982
+
+by Francis G. Howarth and Fred D. Stone, Field No. 22 of F.G. Howarth.
+
+
+
+
+PARATYPES
+: Specimens representing at least 110 individuals from 5
+Hawaiian Islands
+, as listed in appendix 2, including 33 partial to nearly complete skulls, 9 dentaries, 34 associated partial post cranial skeletons, and 60 individual post cranial bones.
+
+
+
+
+TYPE
+LOCALITY:
+Hawaiian Islands
+,
+Maui Island
+, ‘
+Ulupalakua Ranch
+,
+Māhiehie Cave
+,
+
+500 m
+
+,
+20.63°N
+;
+156.39°W
+(WGS 84 datum)
+
+.
+
+
+GEOGRAPHIC AND GEOLOGIC RANGE: Hawaiian Islands, at least the five largest islands, from ca. Middle Pleistocene to Late Holocene (but apparently not into post-1778 historic period), with details as follows. Kaua‘i: infill of sinkhole of Late Pleistocene lithified calcareous dune deposits, and in Early Holocene eolian surface calcareous sand-dune deposits; O‘ahu: in Middle Pleistocene pond deposits of volcanic tuff-cone crater, as well as in later-Pleistoceneto-Holocene composite soil deposits (primarily sedimentary) within limestone sinkholes in emergent Late Pleistocene coral-algal reefs; Moloka‘i: in presumably Polynesian or early postcontact alluvial sediment on floor of a dynamically active piping cave; Maui: on exposed floor, embedded in mineralized crusts on walls and in alluvial sediments of Late Pleistocene or Holocene lava tubes;
+Hawai‘i
+: on exposed floor or in alluvial deposits of Late Pleistocene and Holocene lava tubes; approximate locations of these sites are shown in figure 1, and detailed information on all sites is provided in appendix 4.
+
+
+
+
+FIGURE 1. Map of the main Hawaiian Islands showing relative locations of the principle collecting sites of fossil
+
+Synemporion keana
+
+. See appendix 2 for list of specimens from each site and appendix 4 for detailed descriptions of each site. (
+1
+) Makawehi Point dunes; (
+2)
+Māhā‘ulepū Limestone Sinkhole; (
+3)
+limestone sinkholes near Barbers Point; (
+4)
+Ulupa‘u Head lake deposit; (
+5)
+Lua Lolo Piping Cave
+; (
+6)
+Pu‘u Naio Cave; (
+7)
+Māhiehie Cave; (
+8)
+Pu‘u Mākua Cave; (
+9)
+Ka‘eleku Caverns; (
+10)
+Crystal Cave; (
+11)
+Kahāwaihapapa Cave; (
+12)
+‘Ūmi‘i Manu Cave; (
+13)
+Kahuku Ranch Cave.
+
+
+
+
+ETYMOLOGY: The genus name
+
+Synemporion
+
+root from Greek common noun
+synemporos,
+“fellow traveler or companion,” with addition of the suffix
+-ion
+to form a neuter diminutive, in allusion to the new bat’s former co-occupation of the tectonically mobile Hawaiian Islands with the larger
+
+Lasiurus cinereus semotus
+.
+
+The specific name is a noun in apposition, formed from Hawaiian: the demonstrative
+ke,
+plus
+ana,
+“cave” or “lava tube,” referring to the subterranean provenience of the
+holotype
+and a majority of the
+paratypes
+.
+
+
+
+
+DIAGNOSIS: A small vespertilionid with a dental formula of I1/3, C1/1, P1–2/2, M3/3 = 30–32. Distinguished from other vespertilionid genera with broadly similar dental formulae (
+
+Scotoecus
+,
+Scotozous
+,
+Chalinolobus
+,
+Lasiurus
+,
+Nycticeius, Rhogeesa
+,
+Scotomanes
+,
+Scotophilus
+,
+Otonycteris
+,
+Bauerus
+,
+Pharotis
+,
+Nyctophilus
+
+) by the following traits: upper I2 always absent (sometimes or always present in
+
+Scotozous
+
+and
+
+Chalinolobus
+
+); upper P2 variably present (always absent in
+
+Nycticeius, Rhogeesa
+,
+Scotomanes
+,
+Otonycteris
+,
+Bauerus
+,
+Pharotis
+,
+Nyctophilus
+
+); lower incisors trifid and subequal in size (all bifid in
+
+Otonycteris
+
+and
+
+Scotophilus
+
+; i1 and i2 bifid in most
+Rhogeesa
+, all species of which have i3 reduced to a peg or spicule); M3 crown approximately 1/3 the area of M1 and M2, lacking any trace of a metacone or premetacrista (M3 crown area ½ or more than that of M1 and M2, and premetacrista present, in
+
+Nycticeius
+,
+Nyctophilus
+,
+Scotoecus
+, and
+Scotozous
+
+); m3 with well-developed entocoinid and talonid only slightly narrower than trigoinid (entoconid poorly developed or absent and talonid markedly narrower than trigonid in
+
+Scotomanes
+
+); skull with low rostrum and moderately well-developed forehead that rises abruptly to join with braincase, so that the rostrum profile appears concave in lateral view (rostrum profile flat or convex and forehead break absent in
+
+Lasiurus
+,
+Nycticeius
+,
+Scotomanes
+,
+Scotoecus
+,
+Scotozous
+,
+
+and
+
+Otonycteris
+
+); rostrum relatively narrow (broad in
+
+Scotoecus
+
+,
+
+Scotomanes
+
+, and
+
+Lasiurus
+
+); sagittal crest absent (present in
+
+Lasiurus
+,
+Nycticeius
+,
+Scotozous
+,
+Scotoecus, Rhogeesa
+,
+Scotophilus
+,
+Scotomanes
+,
+Chalinolobus
+,
+Otonycteris
+,
+Bauerus
+,
+Pharotis
+,
+
+and
+
+Nyctophilus
+
+); metacarpal formula III> IV> V (metacarpal formula III = IV = V in
+
+Chalinolobus
+,
+Rhogeessa
+,
+Nycticeius
+,
+Scotoecus
+,
+
+and
+
+Scotomanes
+;
+
+III> IV = V in
+
+Nyctophilus
+
+and
+
+Otonycteris
+
+; III = IV> V in
+
+Scotophilus
+
+).
+
+
+
+
+FIGURE 2. Lateral views of the skull and left dentary of
+A,
+the holotype of
+
+Synemporion keana
+
+(BPBM 159269) compared with
+B,
+
+Lasiurus cinereus semotus
+
+(BPBM 184506). Note the difference in the rostral profile between the two species, including the more robust lower jaw dentition in
+
+Lasiurus
+
+compared with
+
+Synemporion
+
+.
+
+
+
+
+FIGURE 3.
+A,
+Dorsal view and
+B,
+ventral views of the skull of the holotype of
+
+Synemporion keana
+
+(BPBM 159269) compared with
+C,
+dorsal and
+D,
+ventral views of the skull of
+
+Lasiurus cinereus semotus
+
+(BPBM 184506). Note that the rostrum of
+
+Synemporion
+
+is narrower than that of
+
+Lasiurus
+
+, and that
+
+Synemporion
+
+also lacks a sagittal crest.
+
+
+
+
+TABLE 1. Comparative craniodental measurements of Hawaiian bats. All measurements are in mm and were taken as described in the Methods and Materials section. Hypodigm measurements presented as “mean(range)N.” Specimens measured are listed in appendix 3; subadult individuals were excluded from the summaries presented here. The
+
+Synemporion keana
+
+hypodigm measurements include those of the holotype.
+
+
+
+
+
+
+
+Synemporion keana
+
+
+
+
+Lasiurus cinereus semotus
+
+
+
+
+
Holotype
+
Hypodigm
+
Fossil
+
Historic
+
+
+
CRL
+
13.6
+
13.7 (12.9–14.1) 17
+
15.1 (15.1) 1
+
15.3 (14.6–15.9) 14
+
+
+
ZYB
+
9.4
+
9.1 (8.7–9.5) 15
+
–
+
11.5 (11.0–12.4) 16
+
+
+
IOB
+
4.4
+
4.3 (4.1–4.6) 19
+
5.1 (5.1–5.2) 3
+
4.9 (4.7–5.1) 19
+
+
+
BCB
+
7.7
+
7.8 (7.4–8.1) 18
+
8.7 (8.7) 1
+
8.7 (8.2–9.0) 16
+
+
+
PAB
+
5.8
+
5.8 (5.6–6.2) 12
+
8.3 (8.0–8.6) 3
+
7.6 (7.0–8.3) 18
+
+
+
RAB
+
4.8
+
4.8 (4.6–5.0) 23
+
7.5 (6.4–8.5) 4
+
6.8 (6.4–7.2) 19
+
+
+
C–M3
+
4.8
+
4.7 (4.4–4.9) 16
+
5.9 (5.7–6.3) 3
+
5.8 (5.4–6.2) 21
+
+
+
M1–3
+
3.1
+
3.0 (2.7–3.3) 22
+
3.8 (3.6–4.1) 4
+
3.6 (3.4–3.8) 21
+
+
+
MAL
+
9.6
+
9.8 (9.5–10.3) 23
+
12.4 (11.4–13.5) 11
+
11.9 (11.2–12.6) 21
+
+
+
MAH
+
2.8
+
2.9 (2.5–3.2) 26
+
4.4 (3.7–5.2) 14
+
4.0 (3.7–4.7) 21
+
+
+
c–m3
+
5.6
+
5.4 (5.1–5.7) 16
+
6.7 (6.4–7.1) 4
+
6.5 (6.1–6.9) 21
+
+
+
m1–3
+
3.8
+
3.7 (3.4–3.9) 16
+
4.7 (4.3–5.1) 11
+
4.4 (4.1–4.7) 21
+
+
+
m2–3
+
2.4
+
2.4 (2.1–2.6) 22
+
3.0 (2.7–3.3) 19
+
2.8 (2.6–3.1) 21
+
+
+
+
+
+DESCRIPTION: The dental formula of
+
+Synemporion
+
+is I1/3, C1/1, P1–2/2, M3/3 = 30–32, with the small anterior upper premolar (here termed P2 following
+Miller, 1907
+) variably present (figs. 2–5). Among specimens of
+
+Synemporion
+
+with complete palates, P2 is present on both sides in
+4 specimens
+, present on one side and absent on the other side in
+1 specimen
+, and completely absent in
+3 specimens
+including the
+holotype
+(figs. 3, 4). Two half-palates both preserve a P2. Taken together, these observations suggest that P2 was present somewhat more than 50% of the time in
+
+Synemporion
+
+. When present, this tooth is minute and displaced lingually to lie in the angle between the lingual margins of C and P4. Selected craniodental measurements for
+
+S. keana
+
+are given in table 1.
+
+
+
+FIGURE 4. Lateral and occlusal views of the upper dentition of
+A,
+the holotype of
+
+Synemporion keana
+
+(BPBM 159269) compared with that of
+B,
+
+Lasiurus cinereus semotus
+
+(BPBM 184506). Note that a small anterior upper premolar (P2), visible in occlusal view in
+
+Lasiurus
+
+, is absent in the holotype of
+
+Synemporion keana
+
+. This tooth is variably present in
+
+Synemporion
+
+, occurring somewhat more than 50% of the time and sometimes present on one side but absent on the other side in the same individual (see text for discussion).
+
+
+
+
+FIGURE 5. Lateral and occlusal views of the lower dentition of
+A,
+the holotype of
+
+Synemporion keana
+
+(BPBM 159269) compared with that of
+B,
+
+Lasiurus cinereus semotus
+
+(BPBM 184506).
+
+
+
+
+FIGURE 6. Anterior view of the lower incisors and canine of
+A,
+the holotype of
+
+Synemporion keana
+
+(BPBM 159269) compared with those of
+B,
+
+Lasiurus cinereus semotus
+
+(BPBM 184506). The third lower incisor (i3) is absent in the holotype of
+
+Synemporion
+
+but paratypes preserve this tooth, which is always trilobed.
+
+
+
+The single upper incisor of
+
+Synemporion
+
+is roughly two-fifths to one-half height of the upper canine and has about half its crown area, with a small posterointernal cingular cusp (fig. 4). The upper canine is tall, straight, and not recurved (figs. 2, 4). The posterior upper premolar (P4), always present and relatively large, has an unworn crown height about three-fourths that of the canine. An anterolingual cingular cusp is present on P4. M1 and M2 are dilambdodont, subequal in size, and lack a discrete hypocone cusp. Both of these teeth have a postprotocrista that extends from the protocone to the base of metacone, thus closing off the trigon basin posteriorly. The preprotocrista on M1 and M2 extends from protocone to the parastyle rather than to base of paracone. M3 is quite reduced, with a crown area little more than one-third the area of M1 and M2 and it lacks any trace of a metacone or premetacrista. The postparacrista is approximately two-thirds the length of the preparacrista, and the length of the lingual portion of the tooth is greater than the length of the labial portion.
+
+
+The lower incisors of
+
+Synemporion
+
+are mesiodistally broad, trifid; and the crown of i1 substantially overlaps that of i2 when seen in anterior view (figs. 5, 6). The transverse crown width of unworn i1 and i2 about twice diameter of exposed root, and that of unworn i3 (missing in the
+holotype
+although the alveolus is present) is apparently only a little smaller. The lower canine is tall, sharply pointed, and not recurved (figs. 2, 5). Two double-rooted lower premolars are present. The crown length and height of p2 is approximately two-thirds that of p4, which is equal in crown height to m1. Three lower molars are present; m1 and m2 subequal in size, while m3 is somewhat smaller than the anterior molars in all dimensions. The talonid is wider than trigonid in m1 and m2, and both of these teeth are nyctalodont (the postcristid connects hypoconid with hypoconulid, so that talonid basin is narrowly open posteromedially). The last molar (m3) has a talonid that is slightly narrower than trigonid, and lacks a hypoconulid although the hypoconid and entoconid are both well developed.
+
+
+
+TABLE 2. Comparative postcranial measurements of Hawaiian bats. All measurements are in mm and were taken as described in the Methods and Materials section. Hypodigm measurements presented as “mean(range) N.” Specimens measured are listed in appendix 3; all subadult individuals were excluded from the summaries presented here. The
+
+Synemporion keana
+
+hypodigm measurements include those of the holotype.
+
+
+
+
+
+
+
+Synemporion keana
+
+
+
+
+Lasiurus cinereus semotus
+
+
+
+
+
Holotype
+
Hypodigm
+
Fossil
+
Historic
+
+
+
SGW
+
1.2
+
1.3 (1.1–1.4) 25
+
1.8 (1.8–1.9) 5
+
1.7 (1.6–1.9) 19
+
+
+
HUL
+
24.8
+
25.2 (24.0–28.7) 28
+
30.9 (29.4–32.4) 7
+
31.2 (29.4–33.4)18
+
+
+
HPW
+
2.9
+
2.8 (2.5–3.2) 35
+
3.8 (3.7–3.9) 10
+
3.9 (3.6–4.1) 20
+
+
+
HDW
+
2.6
+
2.5 (2.2–2.7) 39
+
3.0 (2.8–3.2) 9
+
2.9 (2.6–3.1) 19
+
+
+
FAL
+
37.5
+
37.9 (35.9–39.7) 39
+
48.0 (46.3–50.6) 6
+
48.7 (45.9–52.7) 32
+
+
+
FAN
+
2.4
+
2.4 (2.1–2.7) 49
+
2.9 (2.6–3.1) 11
+
2.9 (2.6–3.1) 13
+
+
+
FAS
+
1.2
+
1.1 (1.0–1.3) 47
+
1.5 (1.4–1.7) 9
+
1.5 (1.3–1.7) 13
+
+
+
FEL
+
19.0
+
18.8 (17.5–19.9) 20
+
21.4 (21.4)1
+
21.3 (20.4–22.7) 17
+
+
+
FPW
+
1.6
+
1.7 (1.5–1.9) 26
+
2.3 (2.0–2.6) 5
+
2.2 (2.1–2.5) 18
+
+
+
FDW
+
1.5
+
1.5 (1.3–1.6) 24
+
1.8(1.7–1.9)3
+
1.8 (1.7–2.0) 18
+
+
+
TIL
+
17.1
+
16.8 (16.1–18.3) 20
+
20.6 (19.9–21.3) 2
+
20.0 (19.3–21.2) 10
+
+
+
TPW
+
1.6
+
1.6 (1.4–1.7) 23
+
2.0 (1.9–2.1) 2
+
2.0 (1.9–2.1) 13
+
+
+
+
+The skull of
+
+Synemporion
+
+has a low rostrum and a moderately well-developed forehead that rises abruptly to join with braincase, so that the profile of the rostrum appears concave in lateral view (fig. 2). The rostrum is of moderate length and relatively narrow (less than threefourths the width of the braincase) and has a well-developed median sulcus (figs. 2, 3). The frontal portion of braincase is inflated and is higher and slightly wider than the occipital portion. There is no obvious sagittal crest, and the lambdoidal crests are weakly developed and present only laterally. The superior temporal lines are separated about
+1 to 2 mm
+along the sagittal suture. Postorbital processes and supraorbital ridges are entirely absent. The zygomatic arches relatively narrow throughout and lack an expanded dorsal projection. The lachrymal ridge at the anterior rim of orbit is weakly developed.
+
+
+The premaxilla in
+
+Synemporion
+
+lacks a palatal branch and is fused to the maxilla (figs. 2, 3). The narial emargination relatively broad in dorsal view, about 1.5 times as wide as deep, and the anterior palatal emargination even broader, essentially twice as wide as deep. The hard palate extends posteriorly well beyond the tooth row and orbit area. A pair of well-developed but shallow basisphenoid pits are present and are separated by a medial ridge. The cochleae are large, and distance between the cochleae is equivalent to approximately three-fourths the diameter of the cochlea. The dentary is essentially the same as found in most other vespertilionids, with a relatively low coronoid process and a well-developed mental foramen between the roots of C and P2 (fig. 2).
+
+
+Selected postcranial measurements of
+
+Synemporion
+
+are presented in tables 2 and 3. The clavicle is rodlike and lacks any kind of midshaft enlargement or projection. The scapula has a large dorsal articular fossa that is nearly as large as the glenoid fossa, and has an assymetrically bifid coracoid process with the longer branch directed medially. The humerus (figs. 7, 8) has a well-developed greater tuberosity that projects proximally well beyond humeral head; the lesser tuberosity, also well developed, is smaller and does not project beyond head. The bicipital groove along lateral surface of deltoid tuberosity is deep and well defined, and the deltoid tuberosity is high with ridge extending distally for almost one-quarter the total length of humerus. The distal articular surface lies in line with shaft, the trochlea noticeably greater in proximodistal diameter than capitulum, and the medial epicondylar spine projects distally
+0.5 mm
+beyond trochlea. The ulna is fused proximally with radius, and the distalmost ulnar remnant is evident as small, proximally directed, flat spine fused to distal end of lateral border of radius. Metacarpal formula III> IV> V, with metacarpal IV length approximately 94% of that of metacarpal III, and metacarpal V length approximately 87% that of metacarpal III.
+
+
+TABLE 3. Comparative metacarpal measurements and ratios for Hawaiian bats. All measurements are in mm, and ratios are given as percentages.
+
+
+
+
+
Taxon/Specimen
+
II
+
III
+
IV
+
V
+
Ratio IV:III
+
Ratio V:III
+
+
+
+
+Synemporion keana
+
+
+
+
+
BPBM 159265
+
–
+
–
+
40.5
+
37.5
+
–
+
–
+
+
+
BPBM 159267
+
–
+
40.0
+
–
+
35.0
+
–
+
88%
+
+
+
BPBM 1592691
+
39.5
+
42.0
+
–
+
36.5
+
–
+
87%
+
+
+
BPBM 159375
+
–
+
40.0
+
37.5
+
34.5
+
94%
+
86%
+
+
+
BPBM 178052
+
–
+
42.5
+
–
+
34.5
+
–
+
81%
+
+
+
BPBM 178057
+
40.5
+
41.5
+
39.5
+
37.0
+
95%
+
89%
+
+
+
BPBM 178058
+
39.0
+
41.5
+
39.0
+
36.5
+
94%
+
88%
+
+
+
BPBM 178061
+
–
+
42.5
+
39.5
+
36.5
+
93%
+
86%
+
+
+
BPBM 178157
+
40.5
+
42.0
+
39.5
+
36.5
+
94%
+
87%
+
+
+
USNM 498808
+
–
+
41.0
+
38.0
+
35.0
+
93%
+
85%
+
+
+
USNM 520429
+
41.0
+
–
+
40.0
+
37.5
+
–
+
–
+
+
+
USNM 520430
+
–
+
43.5
+
–
+
38.0
+
–
+
87%
+
+
+
USNM 531343
+
40.5
+
43.0
+
39.5
+
37.0
+
92%
+
86%
+
+
+
Mean (x)
+
40.2
+
41.8
+
39.2
+
36.3
+
93.6%
+
86.4%
+
+
+
+Fossil
+
+Lasiurus cinereus semotus
+
+
+
+
+
BPBM 159268
+
51.0
+
51.5
+
46.5
+
41.0
+
90%
+
80%
+
+
+
BPBM 183819
+
–
+
–
+
–
+
40.5
+
–
+
–
+
+
+
BPBM 184971
+
50.5
+
–
+
46.0
+
40.0
+
–
+
–
+
+
+
Mean (x)
+
50.8
+
51.5
+
46.3
+
40.5
+
90%
+
80%
+
+
+
+Historic
+
+Lasiurus cinereus semotus
+
+
+
+
+
BBM 9201
+
54.5
+
56.0
+
51.0
+
43.5
+
91%
+
78%
+
+
+
BBM-X 145165
+
52.0
+
52.5
+
47.5
+
41.0
+
90%
+
78%
+
+
+
BBM-X 145170
+
53.0
+
54.5
+
–
+
42.5
+
–
+
78%
+
+
+
BBM-X 147123
+
54.0
+
55.5
+
49.0
+
44.0
+
88%
+
79%
+
+
+
BBM-X 147125
+
–
+
53.5
+
47.0
+
42.0
+
88%
+
79%
+
+
+
BPBM 161300
+
52.0
+
53.5
+
48.0
+
42.0
+
90%
+
79%
+
+
+
BPBM 175997
+
56.0
+
57.0
+
50.0
+
44.0
+
88%
+
77%
+
+
+
BPBM 179757
+
56.0
+
57.0
+
51.0
+
46.0
+
89%
+
81%
+
+
+
Mean (x)
+
53.9
+
54.9
+
49.1
+
43.1
+
89.1%
+
78.6%
+
+
+
+
+
+1
+Holotype
+
+
+
+
+FIGURE 7.
+A,
+Lateral,
+B,
+anterior,
+C,
+medial, and
+D,
+posterior views of the humeri of the holotype of
+
+Synemporion keana
+
+(on the left in each pair; BPBM 159269) and
+
+Lasiurus cinereus semotus
+
+(on the right; BPBM 184506).
+
+
+
+The vertebral column and pelvis of
+
+Synemporion
+
+are unremarkable and resemble those of other vespertilionids. There is no evidence of vertebral fusion other than in the sacrum. The femur is of usual vespertilionid morphology with the lesser trochanter projecting further proximally than greater trochanter, each distal condyle markedly compressed mediolaterally, and an intercondylar notch that is narrow and deep. The tibia is relatively long and slender, and the fibula is ossified for approximately half of the length of the tibia, the remainder being either threadlike or cartilaginous (not preserved among the material found).
+
+
+
+
\ No newline at end of file
diff --git a/data/7B/05/07/7B0507CD145355E5BE2483370FC43629.xml b/data/7B/05/07/7B0507CD145355E5BE2483370FC43629.xml
new file mode 100644
index 00000000000..8e10db6d80e
--- /dev/null
+++ b/data/7B/05/07/7B0507CD145355E5BE2483370FC43629.xml
@@ -0,0 +1,357 @@
+
+
+
+First contribution to the genera Branchiobaetis and Megabranchiella (Ephemeroptera, Baetidae) in China, with descriptions of two new species
+
+
+
+Author
+
+Tong, Xiaoli
+0000-0003-1731-229X
+Department of Entomology, College of Plant Protection, South China Agricultural University, Guangzhou 510642, Guangdong Province, China
+
+
+
+Author
+
+Zhou, Zhiheng
+https://orcid.org/0009-0009-0763-8713
+Department of Entomology, College of Plant Protection, South China Agricultural University, Guangzhou 510642, Guangdong Province, China
+
+
+
+Author
+
+Wu, Bangyi
+https://orcid.org/0009-0005-6398-5243
+Department of Entomology, College of Plant Protection, South China Agricultural University, Guangzhou 510642, Guangdong Province, China
+
+text
+
+
+ZooKeys
+
+
+2024
+
+2024-10-23
+
+
+1216
+
+
+115
+148
+
+
+
+journal article
+10.3897/zookeys.1216.129803
+2312FE20-8C12-48B2-8D97-CFC920CCF5C2
+
+
+
+
+
+Megabranchiella longusa
+Phlai-ngam & Tungpairojwong, 2022
+
+
+
+
+
+Figs 18
+,
+19
+,
+20
+,
+21
+
+
+
+
+
+
+Megabranchiella longusa
+
+
+: Phlai-ngam & Tungpairojwong in
+
+Phlai-ngam et al. 2022: 16
+
+.
+
+
+
+
+
+
+Material examined.
+
+
+
+One female larva
+on slide;
+Yunnan
+,
+Lushui
+,
+Bajiao River
+(a tributary of the Nujiang River, altitude
+
+1112 m
+
+);
+
+21. iii. 2019
+
+; leg.
+Xiaoli Tong
+
+.
+
+
+
+
+Diagnosis.
+
+
+Female larva (Fig.
+18 a – d
+), body short and flattened, length ~ 4.0 mm; body colour pattern as Fig.
+18 a – c
+.
+
+Head
+.
+
+Labrum nearly semicircular, ~ 1.4 × wider than long; anteromedian notch deep with a small, rounded lobe at base; dorsal surface in distal 1 / 2 with one pair of long, simple setae near midline and irregular row of three medium, simple setae (Fig.
+19 c
+). Left mandible (Fig.
+19 a
+), incisor and kinetodontium fused, incisor with four denticles, kinetodontium with six denticles decreasing in length, prostheca robust, apex with six bluntly denticles and one or two long, spine-like denticles; incisor of right mandible with four denticles, kinetodontium with four denticles, inner margin of innermost denticle with row of small denticles; prostheca robust, apex with comb-like structure, with many denticles apically (Fig.
+19 b
+). Maxilla (Fig.
+19 e
+), galea-lacinia of with three robust canines, base of lacinia with one row of four long, simple setae and one seta perpendicular to lacinia margin; maxillary palp 2 - segmented, apex of segment II with a small cone-shaped projection. Labium (Fig.
+19 d, f
+), glossae shorter and narrower than paraglossae, paraglossae with three rows of long curved setae distoventrally, dorsal surface on distal 1 / 2 with one longitudinal row of two or three long, spine-like setae near inner margin; labial palp 3 - segmented, segment I longer than segments II and III combined, segment II triangular with small protuberance apico-laterally, dorsal surface of segment II with row of two robust, simple setae near distal margin (Fig.
+
+19 g
+
+).
+
+Thorax
+.
+
+Hindwing pads reduced (Fig.
+20 b
+). Forelegs (Fig.
+21 c
+), femur with a row of long, robust, pointed setae along dorsal margin, surface with notched scales anteromedially (Fig.
+20 c
+), villopore present (Fig.
+20 d
+); tibia with a row of long, pointed setae and short, blunt spatulate setae, tibio-patellar suture present; ventral margin of tarsus with one row of four robust, spine-like setae; claws hooked (Fig.
+21 b
+) with one row of 13 acute teeth, subapical setae absent. Middle and hind legs similar to foreleg in structure.
+
+Abdomen
+.
+
+Abdominal tergites and sternites with smooth posterior margins (Fig.
+21 a
+); gills present on abdominal tergites I – VII, gill I oriented ventrally, extremely enlarged and elongated (Fig.
+20 a
+), covering abdominal sternites II –
+VI
+(Fig.
+18 c
+), gills II – VII oriented dorsolaterally, elongated oval similar to tongue blade (Figs
+18 c
+,
+20 a
+), gill margins with long, fine, hair-like setae, ratio of gill length from I – VII = 3.2: 1.7: 1.7: 1.7: 1.5: 1.3: 1.0; paraproct with smooth margin, without marginal spines or spatulate setae, surface with micropores and patch of notch scales (Fig.
+21 d, e
+).
+
+
+
+
+
+
+
+Megabranchiella longusa
+
+a
+gills I, III, V, VII
+b
+hindwing pad
+c
+dorsal margin of femur
+d
+villopore of femur. Red arrow indicates hindwing pad. Red ellipse encloses villopore. Scale bar: 0.1 mm.
+
+
+
+
+
+
+
+
+Megabranchiella longusa
+
+a
+abdominal tergites III – VI
+b
+claw
+c
+foreleg
+d
+notched scales on paraproct surface
+e
+paraproct.
+
+
+
+
+
+Distribution
+
+
+
+(Fig.
+23
+).
+
+Thailand
+(
+Chiang Mai
+and
+Nan
+Provinces) and
+China
+(
+Yunnan Province
+).
+
+
+
+
+Larval habitat
+
+
+
+(Fig.
+22 c
+).
+
+The species was collected in a swift, unshaded stream with cobble substrate at an altitude of ~
+1100 m
+in
+Yunnan
+,
+China
+.
+
+
+
+
+
+
+Representative sites of larval habitat
+a
+
+Branchiobaetis megasinus
+
+sp. nov.
+Upper reaches of Liuxihe River, Conghua, Guangdong
+b
+
+Branchiobaetis borealis
+
+sp. nov.
+Lapu River, Weixi, Yunnan
+c
+
+Megabranchiella longusa
+
+Phlai-ngam & Tungpairojwong. Bajiao River, Lushui, Yunnan.
+
+
+
+
+
+
+
+Distribution map of the genera
+
+Branchiobaetis
+
+and
+
+Megabranchiella
+
+in China.
+
+
+
+
+
+Remarks.
+
+
+Geographically, this record represents the farthest distribution north of the genus
+
+Megabranchiella
+
+so far. We expect that more species of the genus will be discovered with the expansion of the investigation range in
+China
+.
+
+
+
+
\ No newline at end of file